A new statistical method for estimating divergence dates of species from DNA sequence data by a molecular clock approach is developed. This method takes into account effectively the information contained in a set of DNA sequence data. The molecular clock of mitochondrial DNA (mtDNA) was calibrated by setting the date of divergence between primates and ungulates at the Cretaceous-Tertiary boundary (65 million years ago), when the extinction of dinosaurs occurred. A generalized least-squares method was applied in fitting a model to mtDNA sequence data, and the clock gave dates of 92.3 +/- 11.7, 13.3 +/- 1.5, 10.9 +/- 1.2, 3.7 +/- 0.6, and 2.7 +/- 0.6 million years ago (where the second of each pair of numbers is the standard deviation) for the separation of mouse, gibbon, orangutan, gorilla, and chimpanzee, respectively, from the line leading to humans. Although there is some uncertainty in the clock, this dating may pose a problem for the widely believed hypothesis that the pipedal creature Australopithecus afarensis, which lived some 3.7 million years ago at Laetoli in Tanzania and at Hadar in Ethiopia, was ancestral to man and evolved after the human-ape splitting. Another likelier possibility is that mtDNA was transferred through hybridization between a proto-human and a proto-chimpanzee after the former had developed bipedalism.

Dating of the human-ape splitting by a molecular clock of mitochondrial DNA.

The Brush Foundation studies on human growth and development, begun in 1931 and terminated in 1942, have been intensively reviewed and studied by Dr Greulich and Miss Pyle in the formulation of this Radiographic Atlas of Skeletal Development of the Hand and Wrist Serial radiographs of from 2 to 20 hand-films made at successive examinations of each of 1000 boys and girls made up the radiographic material Standards were selected that were judged to be the most representative of the central tendency or anatomic mode of each chronologic age group from birth through 18 years

Radiographic Atlas of Skeletal Development of the Hand and Wrist

Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.

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Endurance running and the evolution of homo

Nonhuman primates, our closest biological relatives, play important roles in the livelihoods, cultures, and religions of many societies and offer unique insights into human evolution, biology, behavior, and the threat of emerging diseases. They are an essential component of tropical biodiversity, contributing to forest regeneration and ecosystem health. Current information shows the existence of 504 species in 79 genera distributed in the Neotropics, mainland Africa, Madagascar, and Asia. Alarmingly, ~60% of primate species are now threatened with extinction and ~75% have declining populations. This situation is the result of escalating anthropogenic pressures on primates and their habitats—mainly global and local market demands, leading to extensive habitat loss through the expansion of industrial agriculture, large-scale cattle ranching, logging, oil and gas drilling, mining, dam building, and the construction of new road networks in primate range regions. Other important drivers are increased bushmeat hunting and the illegal trade of primates as pets and primate body parts, along with emerging threats, such as climate change and anthroponotic diseases. Often, these pressures act in synergy, exacerbating primate population declines. Given that primate range regions overlap extensively with a large, and rapidly growing, human population characterized by high levels of poverty, global attention is needed immediately to reverse the looming risk of primate extinctions and to attend to local human needs in sustainable ways. Raising global scientific and public awareness of the plight of the world’s primates and the costs of their loss to ecosystem health and human society is imperative.

/pdf/impending-extinction-crisis-of-the-world-s-primates-why-32dorf7ucr.pdf

Impending extinction crisis of the world's primates: why primates matter

"Wolff's law" is a concept that has sometimes been misrepresented, and frequently misunderstood, in the anthropological literature. Although it was originally formulated in a strict mathematical sense that has since been discredited, the more general concept of "bone functional adaptation" to mechanical loading (a designation that should probably replace "Wolff's law") is supported by much experimental and observational data. Objections raised to earlier studies of bone functional adaptation have largely been addressed by more recent and better-controlled studies. While the bone morphological response to mechanical strains is reduced in adults relative to juveniles, claims that adult morphology reflects only juvenile loadings are greatly exaggerated. Similarly, while there are important genetic influences on bone development and on the nature of bone's response to mechanical loading, variations in loadings themselves are equally if not more important in determining variations in morphology, especially in comparisons between closely related individuals or species. The correspondence between bone strain patterns and bone structure is variable, depending on skeletal location and the general mechanical environment (e.g., distal vs. proximal limb elements, cursorial vs. noncursorial animals), so that mechanical/behavioral inferences based on structure alone should be limited to corresponding skeletal regions and animals with similar basic mechanical designs. Within such comparisons, traditional geometric parameters (such as second moments of area and section moduli) still give the best available estimates of in vivo mechanical competence. Thus, when employed with appropriate caution, these features may be used to reconstruct mechanical loadings and behavioral differences within and between past populations.

Who's afraid of the big bad Wolff?: "Wolff's law" and bone functional adaptation.

Body proportions, skeletal allometry and locomotion in the hadar hominids: a reply to Wolpoff

Current views on the function of the deltoid and rotator cuff muscles emphasize their roles in arm-raising as participants in a scapulohumeral force “couple.” The acceptance of such a mechanism is based primarily on a 1944 EMG study of human shoulder muscle action. More recently, it has been suggested that shoulder joint stabilization constitutes a second and equally important function of the cuff musculature, especially in nonhuman primates which habitually use their forelimbs in overhead postural and locomotor activities. Few comparative data exist, however, on the actual recruitment patterns of these muscles in different species. In order to assess the general applicability of a scapulohumeral force couple model, and the functional significance of the differential development of the scapulohumeral musculature among primate species, we have undertaken a detailed study of shoulder muscle activity patterns in nonhuman primates employing telemetered electromyography, which permits examination of unfettered natural behaviors and locomotion. The results of our research on the chimpanzee, Pan troglodytes, on voluntary reaching and two forms of “arboreal” locomotion reveal four ways in which previous perceptions of the function of the scapulohumeral muscles must be revised: (1) the posterior deltoid is completely different in function from the middle and anterior regions of this muscle; (2) the integrity of the glenohumeral joint during suspensory postures is not maintained solely by osseoligamentous structures; (3) the function of teres minor is entirely different from that of the other rotator cuff muscles and is more similar to the posterior deltoid and teres major; and (4) each remaining member of the rotator cuff plays a distinct, and often unique, role during natural behaviors. These results clearly refute the view that the muscles of the rotator cuff act as a single functional unit in any way, and an alternative to the force couple model is proposed.

EMG of scapulohumeral muscles in the chimpanzee during reaching and “arboreal” locomotion

The importance of knuckle-walking in the locomotor repertoire of African apes raises the possibility that the long digital flexors may be specially adapted more to meet the demands of ground quadrupedalism than those of suspension. To investigate this possibility, the activities of the flexor digitorum superficialis and flexor digitorum profundus were studied by means of telemetered electromyography in three chimpanzees. Results clearly indicate that the fasciculi of these muscles to digits bearing weight in knuckle-walking are not called upon to contract in quadrupedal postures or in slow and moderately fast quadrupedal locomotion except to help clear the fingers from the ground as the forelimb begins its recovery stroke. At the most rapid speeds, a slight to moderate level of activity sometimes occurs in the latter half of stance phase. The long digital flexors display maximum and sustained activity during suspension. It is concluded that any role for these muscles in maintenance of stability at the metacarpophalangeal joints during knuckle-walking must be predominantly passive. Prominent markings for insertions of these muscles in a fossil hand (such as O.H. 7) suggest use of the forelimb in suspensory climbing behaviors.

Telemetered electromyography of flexor digitorum profundus and flexor digitorum superficialis in Pan troglodytes and implications for interpretation of the O. H. 7 hand.

By most accounts, the upper limb of the chimpanzee is primarily adapted to suspensory postures and locomotion. In order to determine how the derived morphology of the chimpanzee forelimb has affected the form of quadrupedal locomotion displayed by these animals, electromyographic activity patterns of 10 shoulder muscles during knuckle-walking in two chimpanzee subjects were analysed and compared to data on the opossum and cat taken from the literature. Telemetered electromyography coupled with simultaneous video recording was employed in order to study unfettered locomotion in the chimpanzee subjects.

Chimpanzees are characterized by a quadrupedal gait in which the hind limb overstrides the ipsilateral forelimb. Forelimb position in the plane of abduction/adduction is significantly affected by whether the hind limb passes inside or outside its ipsilateral forelimb. The degree of abduction adduction of the forelimb, in turn, influences many of the muscle activity patterns. That is, some muscles would be more frequently or less frequently active, depending on whether the arm was relatively abducted or adducted during a stride. Thus, there can be no single motor programme that generates the step cycle in chimpanzees.

While there are some parallels between muscle recruitment patterns for chimpanzee, opossum and cat quadrupedalism, the results of this study also indicate that many aspects of muscle use in chimpanzees have been significantly influenced by factors related to increased mobility of the upper limb. Finally, this study has revealed that moving the arm forward during swing phase of knuckle-walking is not a simple product of muscular elTort. and that other mechanisms must be involved. However, it is unclear at present exactly what these mechanisms may be.

EMG of chimpanzee shoulder muscles during knuckle‐walking: problems of terrestrial locomotion in a suspensory adapted primate

Theories about the functions of the foot muscles have centered on their role in arch support. Previous anatomical and electromyographic studies (reviewed herein) have demonstrated that the arches are normally maintained by bones and ligaments. This study reports an electromyographic investigation of five foot muscles (flexor digitorum longus, flexor digitorum brevis, flexor accessorius, abductor hallucis, and abductor digiti quinti) conducted on four humans. The three toe flexors act together to resist extension of the toes during the stance phase of locomotion. Despite the large flexor accessorius in humans, neither this muscle nor the flexor digitorum brevis are preferentially recruited over the flexor digitorum longus for any normal posture or locomotion. The abductors affect the mediolateral distribution of pressure by positioning the forefoot. We suggest that the foot muscles play an important role in positioning of the forces on the foot in both posture and locomotion. Future electromyographic experiments on human and ape foot muscles in conjunction with detailed studies of early hominid fossils promise to elucidate the pathways of human locomotor evolution.

Jack T. Stern

Papers

Body proportions, skeletal allometry and locomotion in the hadar hominids: a reply to Wolpoff

EMG of scapulohumeral muscles in the chimpanzee during reaching and “arboreal” locomotion

Telemetered electromyography of flexor digitorum profundus and flexor digitorum superficialis in Pan troglodytes and implications for interpretation of the O. H. 7 hand.

EMG of chimpanzee shoulder muscles during knuckle‐walking: problems of terrestrial locomotion in a suspensory adapted primate

Electromyographic studies of the human foot: experimental approaches to hominid evolution.