Body size prediction from juvenile skeletal remains

There are currently no methods for predicting body mass from juvenile skeletal remains and only a very limited number for predicting stature. In this study, stature and body mass prediction equations are generated for each year from 1 to 17 years of age using a subset of the Denver Growth Study sample, followed longitudinally (n = 20 individuals, 340 observations). Radiographic measurements of femoral distal metaphyseal and head breadth are used to predict body mass and long bone lengths are used to predict stature. In addition, pelvic bi-iliac breadth and long bone lengths are used to predict body mass in older adolescents. Relative prediction errors are equal to or smaller than those associated with similar adult estimation formulae. Body proportions change continuously throughout growth, necessitating age-specific formulae. Adult formulae overestimate stature and body mass in younger juveniles, but work well in 17-year-olds from the sample, indicating that in terms of body proportions they are representative of the general population. To illustrate use of the techniques, they are applied to the juvenile Homo erectus (ergaster) KNM-WT 15000 skeleton. New body mass and stature estimates for this specimen are similar to previous estimates derived using other methods. Body mass estimates range from 50 to 53 kg, and stature was probably slightly under 157 cm, although a precise stature estimate is difficult to determine due to differences in linear body proportions between KNM-WT 15000 and the Denver reference sample.     

Just how strapping was KNM-WT 15000?

For over twenty years, the young, male Homo erectus specimen KNM-WT 15000 has been the focus of studies on growth and development, locomotion, size, sexual dimorphism, skeletal morphology, and encephalization, often serving as the standard for his species. Prior research on KNM-WT 15000 operates under the assumption that H. erectus experienced a modern human life history, including an adolescent growth spurt. However, recent fossil discoveries, improvements in research methods, and new insights into modern human ontogeny suggest that this may not have been the case. In this study, we examine alternative life history trajectories in H. erectus to re-evaluate adult stature estimates for KNM-WT 15000. We constructed a series of hypothetical growth curves by modifying known human and chimpanzee curves, calculating intermediate growth velocities, and shifting the age of onset and completion of growth in stature. We recalculated adult stature for KNM-WT 15000 by increasing stature at death by the percentage of growth remaining in each curve. The curve that most closely matches the life history events experienced by KNM-WT 15000 prior to death indicates that growth in this specimen would have been completed by 12.3 years of age. These results suggest that KNM-WT 15000 would have experienced a growth spurt that had a lower peak velocity and shorter duration than the adolescent growth spurt in modern humans. As a result, it is likely that KNM-WT 15000 would have only attained an adult stature of 163 cm (∼5′4″), not 185 cm (∼6′1″) as previously reported. KNM-WT 15000's smaller stature has important implications for evolutionary scenarios involving early genus Homo.     

Skeletal age, dental age, and the maturation of KNM-WT 15000

The skeleton of the Homo erectus boy from West Lake Turkana, Kenya (KNM-WT 15000), is remarkably complete, and this individual has thus provided a case study for several researchers examining Homo erectus growth. Using data from a longitudinal study of Montreal French-Canadian children, it is shown that while dental and skeletal ages match reasonably well at the level of a sample of children, individuals can display differences between skeletal and dental ages of 2 years or more. Furthermore, the relationship between these two markers may change over time in individual children. It is also possible to find children with patterns of dental maturation similar to KNM-WT 15000's pattern in the Montreal sample. Therefore, neither the discrepancy between skeletal age and dental age alone nor the pattern of dental maturation as assessed by dental stages precludes a human-like pattern of growth, including an adolescent growth spurt, for this individual. Some indicators (e.g., estimated body size for predicted age, and enamel formation) do suggest possible growth-patterning differences from modern humans, and therefore earlier maturation is a reasonable hypothesis, but caution is warranted, given the large degree of modern human variation in developmental markers and the inherent uncertainty in precise estimation of KNM-WT 15000's maturational parameters.     

Vertebrae numbers of the early hominid lumbar spine

General doctrine holds that early hominids possessed a long lumbar spine with six segments. This is mainly based on Robinson's (1972) interpretation of a single partial Australopithecus africanus skeleton, Sts 14, from Sterkfontein, South Africa. As its sixth last presacral vertebra exhibits both thoracic and lumbar characteristics, current definitions of lumbar vertebrae and lumbar ribs are discussed in the present study. A re-analysis of its entire preserved vertebral column and comparison with Stw 431, another partial A. africanus skeleton from Sterkfontein, and the Homo erectus skeleton KNM-WT 15000 from Nariokotome, Kenya, did not provide strong evidence for the presence of six lumbar vertebrae in either of these early hominids. Thus, in Sts 14 the sixth last presacral vertebra has on one side a movable rib. In Stw 431, the corresponding vertebra shows indications for a rib facet. In KNM-WT, 15000 the same element is very fragmentary, but the neighbouring vertebrae do not support the view that it is L1. Although in all three fossils the transitional vertebra at which the articular facets change orientation seems to be at Th11, this is equal to a large percentage of modern humans. Indeed, a modal number of five lumbar vertebrae, as in modern humans, is more compatible with evolutionary principles. For example, six lumbar vertebrae would require repetitive shortening and lengthening not only of the lumbar, but also of the entire precaudal spine. Furthermore, six lumbar vertebrae are claimed to be biomechanically advantageous for early hominid bipedalism, yet an explanation is lacking as to why the lumbar region should have shortened in later humans. All this raises doubts about previous conclusions for the presence of six lumbar vertebrae in early hominids. The most parsimonious explanation is that they did not differ from modern humans in the segmentation of the vertebral column.     

Comparison of inverse-dynamics musculo-skeletal models of AL 288-1 Australopithecus afarensis and KNM-WT 15000 Homo ergaster to modern humans, with implications for the evolution of bipedalism

Size and proportions of the postcranial skeleton differ markedly between Australopithecus afarensis and Homo ergaster, and between the latter and modern Homo sapiens. This study uses computer simulations of gait in models derived from the best-known skeletons of these species (AL 288-1, Australopithecus afarensis, 3.18 million year ago) and KNM-WT 15000 (Homo ergaster, 1.5-1.8 million year ago) compared to models of adult human males and females, to estimate the required muscle power during bipedal walking, and to compare this with those in modern humans. Skeletal measurements were carried out on a cast of KNM-WT 15000, but for AL 288-1 were taken from the literature. Muscle attachments were applied to the models based on their position relative to the bone in modern humans. Joint motions and moments from experiments on human walking were input into the models to calculate muscle stress and power. The models were tested in erect walking and 'bent-hip bent-knee' gait. Calculated muscle forces were verified against EMG activity phases from experimental data, with reference to reasonable activation/force delays. Calculated muscle powers are reasonably comparable to experimentally derived metabolic values from the literature, given likely values for muscle efficiency. The results show that: 1) if evaluated by the power expenditure per unit of mass (W/kg) in walking, AL 288-1 and KNM-WT 15000 would need similar power to modern humans; however, 2) with distance-specific parameters as the criteria, AL 288-1 would require to expend relatively more muscle power (W/kg.m(-1)) in comparison to modern humans. The results imply that in the evolution of bipedalism, body proportions, for example those of KNM-WT 15000, may have evolved to obtain an effective application of muscle power to bipedal walking over a long distance, or at high speed.     

Homo floresiensis and the evolution of the hominin shoulder

The holotype of Homo floresiensis, diminutive hominins with tiny brains living until 12,000 years ago on the island of Flores, is a partial skeleton (LB1) that includes a partial clavicle (LB1/5) and a nearly complete right humerus (LB1/50). Although the humerus appears fairly modern in most regards, it is remarkable in displaying only 110 degrees of humeral torsion, well below modern human average values. Assuming a modern human shoulder configuration, such a low degree of humeral torsion would result in a lateral set to the elbow. Such an elbow joint would function more nearly in a frontal than in a sagittal plane, and this is certainly not what anyone would have predicted for a tool-making Pleistocene hominin. We argue that Homo floresiensis probably did not have a modern human shoulder configuration: the clavicle was relatively short, and we suggest that the scapula was more protracted, resulting in a glenoid fossa that faced anteriorly rather than laterally. A posteriorly directed humeral head was therefore appropriate for maintaining a normally functioning elbow joint. Similar morphology in the Homo erectus Nariokotome boy (KNM-WT 15000) suggests that this shoulder configuration may represent a transitional stage in pectoral girdle evolution in the human lineage.     

The place of Neandertals in the evolution of hominid patterns of growth and development

This study uses the two developmental fields of dental maturation and femoral growth to determine if the pattern of growth and development in Neandertals (archaic Homo sapiens) was intermediate between that of Homo erectus and recent modern humans. Specimens used in the analysis included Neandertals and Upper Palaeolithic early modern Homo sapiens from Europe and individuals from two recent modern human populations. Ontogenetic data for the H. erectus adolescent KNM-WT 15000 and for Gorilla gorilla were included for comparison. Previous reports have indicated that H. erectus demonstrates a pattern of ontogeny characterized by earlier and more rapid linear growth than in modern humans. Results reported here demonstrate that Upper Paleolithic early modern Homo sapiens display a growth trajectory indistinguishable from that of recent modern humans. The pattern of Neandertal ontogeny is not intermediate between the pattern displayed in H. erectus and the derived pattern seen in the modern reference samples and the early modern H. sapiens sample. The Neandertal growth trajectory is consistent with either slow linear growth or advanced dental development.     

Estimation of body size and physique from hominin skeletal remains

Three methods of measuring stature from skeletal remains are reviewed: the reconstructed skeletal length, the correspondence of long bone length to stature and the regression of stature on long bone length. Each involves problems and difficulties. For the anthropologist, there is the additional problem of applying findings from extant taxa to extinct taxa with potentially different morphologies and limb proportions. Of the various studies involving regression of the stature the findings of Trotter and Gleser are judged the most robust and useful notwithstanding problems and limitations. The lumbar vertebrae are potentially important as stature predictors. Estimation of body mass from the skeleton is also beset with problems. Eight methods are reviewed: Hartwig-Scherer's taxon independent solution, four methods involving measurements from the weight-bearing appendicular skeleton, Ruff's method using the length of the reconstructed skeleton and an estimate of body breadth, estimates from the total skeletal mass and estimates from the body mass index when the stature is known approximately. Lumbar vertebrae provide reasonable estimates of both body mass and stature and thus by derivation the body mass index. At present both forensic scientists and anthropologists lack adequate data and methods to estimate body size and shape from hominin skeletons. A further large and well-designed study using magnetic resonance imaging is required.     

萨拉乌苏河流域化石智人的身高

采用股骨骨干下部横径等,根据中国汉族男女性长骨碎骨的身高推断回归方程推测有比较明确地层记载的萨拉乌苏河流域化石智人的身高。据此计算复原股骨长。采用一系列身高推测公式,得出该标本可能的身高范围并比较。该化石的身材比北京直立人高。与晚更新世智人相比,该流域化石智人的身材比山顶洞化石人类矮,比柳江化石人类高。     

Estimating stature in fossil hominids: which regression model and reference sample to use?

coResearchers have long appreciated the significant relationship between body size and an animal's overall adaptive strategy and life history. However, much more emphasis has been placed on interpreting body size than on the actual calculation of it. One measure of size that is especially important for human evolutionary studies is stature. Despite a long history of investigation, stature estimation remains plagued by two methodological problems: (1) the choice of the statistical estimator, and (2) the choice of the reference population from which to derive the parameters.This work addresses both of these problems in estimating stature for fossil hominids, with special reference to A.L. 288-1 (Australopithecus afarensis) and WT 15000 (Homo erectus). Three reference samples of known stature with maximum humerus and femur lengths are used in this study: a large (n=2209) human sample from North America, a smaller sample of modern human pygmies (n=19) from Africa, and a sample of wild-collected African great apes (n=85). Five regression techniques are used to estimate stature in the fossil hominids using both univariate and multivariate parameters derived from the reference samples: classical calibration, inverse calibration, major axis, reduced major axis and the zero-intercept ratio model. We also explore a new diagnostic to test extrapolation and allometric differences with multivariate data, and we calculate 95% confidence intervals to examine the range of variation in estimates for A.L. 288-1, WT 15000 and the new Bouri hominid (contemporary with [corrected] Australopithecus garhi). Results frequently vary depending on whether the data are univariate or multivariate. Unique limb proportions and fragmented remains complicate the choice of estimator. We are usually left in the end with the classical calibrator as the best choice. It is the maximum likelihood estimator that performs best overall, especially in scenarios where extrapolation occurs away from the mean of the reference sample. The new diagnostic appears to be a quick and efficient way to determine at the outset whether extrapolation exists in size and/or shape of the long bones between the reference sample and the target specimen.     

Relative limb strength and locomotion in Homo habilis

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.     

Femur/stature ratio and estimates of stature in children.

The present study examines the relationship between femur length and stature in children between the ages of 8 and 18 years. In previous investigations, my colleagues and I reported the surprising finding that femur length bears a nearly constant relationship to stature in adult humans regardless of ethnicity or gender. This earlier study revealed that the femur/stature ratio averages 26.74% in adult humans, and that using the ratio to predict stature from femur length yields remarkably accurate estimates. The current study shows that femur/stature ratios of children between the ages of 8 and 11 differ significantly from their older counterparts. Between the ages of 12 and 18, there are no significant differences due to age in the femur/stature ratio; however, there are significant differences in this age group attributable to gender. This study also shows that the worldwide average adult femur/stature ratio does not adequately describe children in this age range. This study strongly documents the adolescent growth spurt in the femur/stature ratios of both males and females at the precise time one would expect to see the spurt occur (10-12 in females; 12-14 in males). This growth follows a nearly identical trajectory in both genders, with relative femur growth dominating before the peak years of the growth spurt, and relative stature growth dominating afterward. This accounts for the ratio's rise to maximum values just before peak growth, and its decline toward the adult ratio thereafter. These findings require us to use separate adolescent femur/stature ratios of 27.16 (females) and 27.44 (males) to estimate the stature of children between the ages of 12 and 18. Preliminary testing shows these ratios to be more accurate in estimating stature than the properly selected Trotter and Gleser adult regression equation. Use of the adolescent male ratio with the Homo erectus juvenile WT 15000 results in a lower stature estimate (157.4 cm) than previously reported. It is suggested that continued testing of the ratio occur, but that the values herein derived may be useful in routine forensic cases involving children in this age range, and with subadult paleontological specimens.     

Femoral lengths and stature in Plio-Pleistocene hominids

This study reports the femoral lengths of 31 Plio-Pleistocene hominids dated between 3.1 and 0.7 million years ago, and uses those lengths to estimate stature by way of the femur-stature ratio reported by Feldesman et al. (Am. J. Phys. Anthropol. 78:219-220, 1989). By this method the average female Australopithecus afarensis is 105 cm and the average male is 151 cm. The respective values are 115 and 138 cm for A. africanus. As defined by Howell (In VJ Maglio and HBS Cooke (eds): The Evolution of African Mammals. Cambridge: Harvard University Press, 1978) and Johanson et al. (Kirtlandia 28:1-14, 1978), Homo habilis is a sexually dimorphic species, with females standing 118 cm and males 157 cm. Such apparently strong dimorphism may be due to the possibility that there are actually two species of nonrobust hominids between 2 and 1.7 m.y.a. The estimate for the female Australopithecus boisei is 124 cm and for the male, 137 cm, but these estimates are especially difficult to be certain of because there are no femora that can be positively identified as male A. boisei. Australopithecus robustus is estimated to be 110 cm (female) and 132 cm (male). African Homo erectus stood 160 cm (female) and 180 cm (male). From these estimates several generalizations are apparent. First, there is apparently strong sexual dimorphism in stature in A. afarensis and H. habilis, but less in the other species. Second, the "robust" australopithecines were relatively small statured. Third, it is apparently not true that humans have been getting progressively taller throughout their evolutionary history. Some individuals were as tall as modern humans 3 m.y.a., by 2 m.y.a. one individual stood about 173 cm, and by 1.7 m.y.a. a stature of 180+ cm was not uncommon.     

Femoral/humeral strength in early African Homo erectus

Lower-to-upper limb-bone proportions give valuable clues to locomotor behavior in fossil taxa. However, to date only external linear dimensions have been included in such analyses of early hominins. In this study, cross-sectional measures of femoral and humeral diaphyseal strength are determined for the two most complete early Homo erectus (or ergaster) associated skeletons--the juvenile KNM-WT 15000 and the adult KNM-ER 1808. Modern comparative samples include an adult human skeletal sample representative of diverse body shapes, a human longitudinal growth series, and an adult chimpanzee sample. When compared to appropriately age-matched samples, both H. erectus specimens fall very close to modern human mean proportions and far from chimpanzee proportions (which do not overlap with those of humans). This implies very similar mechanical load-sharing between the lower and upper limbs, and by implication, similar locomotor behavior in early H. erectus and modern humans. Thus, by the earliest Pleistocene (1.7 Ma), completely modern patterns of bipedal behavior were fully established in at least one early hominin taxon.     

Gamma-ray spectrometric dating of late Homo erectus skulls from Ngandong and Sambungmacan, Central Java, Indonesia

Hominid fossils from Ngandong and Sambungmacan, Central Java, Indonesia, are considered to be the most anatomically derived and youngest representatives of Homo erectus. Nondestructive gamma-ray spectrometric dating of three of these Homo erectus skulls showed that all samples underwent uranium leaching. Nevertheless, we could establish minimum age estimates of around 40ka, with an upper age limit of around 60 to 70ka. This means that the Homo erectus of Java very likely survived the Toba eruption and may have been contemporaneous with the earliest Homo sapiens in Southeast Asia and Australasia.     

No skeletal dysplasia in the nariokotome boy KNM‐WT 15000 (homo erectus)—A reassessment of congenital pathologies of the vertebral column

The Nariokotome boy skeleton KNM‐WT 15000 is the most complete Homo erectus fossil and therefore is key for understanding human evolution. Nevertheless, since Latimer and Ohman (2001) reported on severe congenital pathology in KNM‐WT 15000, it is questionable whether this skeleton can still be used as reference for Homo erectus skeletal biology. The asserted pathologies include platyspondylic and diminutive vertebrae implying a disproportionately short stature; spina bifida; condylus tertius; spinal stenosis; and scoliosis. Based on this symptom complex, the differential diagnosis of spondyloepiphyseal dysplasia tarda, an extremely rare form of skeletal dysplasia, has been proposed. Yet, our reanalysis of these pathologies shows that the shape of the KNM‐WT 15000 vertebrae matches that of normal modern human adolescents. The vertebrae are not abnormally flat, show no endplate irregularities, and thus are not platyspondylic. As this is the hallmark of spondyloepiphyseal dysplasia tarda and related forms of skeletal dysplasia, the absence of platyspondyly refutes axial dysplasia and disproportionate dwarfism. Furthermore, we neither found evidence for spina bifida occulta nor manifesta, whereas the condylus tertius, a developmental anomaly of the cranial base, is not related to skeletal dysplasias. Other fossils indicate that the relatively small size of the vertebrae and the narrow spinal canal are characteristics of early hominins rather than congenital pathologies. Except for the recently described signs of traumatic lumbar disc herniation, the Nariokotome boy fossil therefore seems to belong to a normal Homo erectus youth without pathologies of the axial skeleton. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

A new brain endocast of Homo erectus from Hulu Cave, Nanjing, China

A new brain endocast of Homo erectus from Hulu Cave, Tangshan, Nanjing is described and compared with a broad sample of endocasts of H. erectus, Neanderthals, and recent modern humans. The Nanjing 1 endocast is reconstructed based on two portions of endocranial casts taken from the original fossil fragments. The fossil was discovered in 1993, near Nanjing, South China and is dated to ～ 0.58-0.62 Ma. The cranial capacity is ～ 876 cc, as determined by endocast water displacement. There are some common features of Nanjing 1 and other H. erectus endocasts that differentiate them from the Neanderthals and modern humans in our sample. These include small cranial capacity, low height dimensions, simple middle meningeal vessel patterns, a high degree of cerebral-over-cerebellar lobe overhang, elongated and quite separated cerebellar lobes, and a narrow, low, short and flat frontal region. Some features are found to vary among H. erectus, Neanderthals and modern humans, such as the lateral Sylvian fissure position and the venous sinus and petalial patterns. The Nanjing 1 endocast has unique, large, superior frontal convolutions, and strongly protruding Broca's caps. In contrast to other Chinese H. erectus from Hexian and Zhoukoudian, Nanjing 1 lacks strong posterior projection of the occipital lobes. Bivariate and principal component analyses indicate that the small volume and shape of Nanjing 1 is most similar to KNM-WT 15000, KNM-ER 3883, Sangiran 2 and Hexian, illustrating the combination of narrow, low, and short frontal lobes with wide posterior lobes.     

The subspecies ofHomo erectus

ClassifyingHomo erectus into subspecies can be based on either temporal or geographical differences, but there is no accepted system for using both. This can be done with subspecies names consisting of two elements — a prefix ofneo, meso, orpaleo to indicate grade, followed by a geographical term ofeuropus, africus, sinicus, orindicus to indicate line. Thus Rhodesian isHomo erectus neoafricus, Ngandong isHomo erectus neoindicus, Peking isHomo erectus mesosinicus, ER 3733 isHomo erectus paleoafricus, etc.     

Estimation of stature from the skeletal reconstruction of an immature Neandertal from Dederiyeh cave, Syria

Skeletal reconstruction of a child Neandertal unearthed at Dederiyeh Cave, Syria in 1993, is undertaken and the acquired stature discussed. Although the skeletal remains were well preserved, the reconstruction required several assumptions to be made because of the immature status of the specimen. The assumptions were mainly concerned with distances between bones in the inter-vertebral spaces and in the joints of the hip, knee, and ankle. These were estimated from X-ray films of modern children and data from previous studies. Stature was directly measured on the reconstruction, and found to be 79.2 cm. After corrections for soft tissue thickness and shrinkage of the casts, the stature became 81.7 cm. This estimate is consistent with estimates based on regression equations of long bone lengths, especially from those of the lower extremity. In comparison with longitudinal data for white American boys, the assessment of stature for Dederiyeh varied according to the estimated age. For a younger estimated age, the stature falls in the lower half of the white American range of variation, but with an older estimated age, it falls below the lower limit of the range of variation. Other immature Neandertals including two European specimens, Roc de Marsal and La Ferrassie 6, fall below the lower limit of the 5th to 95th percentile range based on the estimated statures from their long bone lengths. More comprehensive age assessment covering both fossil and modern humans is required before accurate conclusions in relation to Neandertal growth can be drawn.