The capitular joint of the first rib in primates: a re-evaluation of the proposed link to locomotion

Evidence exists that in Australopithecus afarensis the head of the first rib articulated with the body of the first thoracic vertebra but not with the body of the seventh cervical vertebra (Ohman, 1983, 1986). Thus, the Hadar hominid would have differed from most primates, in which both these vertebrae are involved in formation of the first costal capitular joint. Indeed, Ohman (1986) has claimed that a univertebral pattern is unique to modern and fossil hominids among primates. He offered various theories on the adaptive significance of this trait, chief among which was a link to freeing the upper limb from any role in locomotion. Believing that Ohman's statement about the distribution of the univertebral pattern in living forms was based on inadequate samples, we have compiled data on the first costal capitular joint in a wider range of primate genera. Our observations demonstrate that the univertebral pattern, rather than being unique to hominids, is common among siamangs, occurs in an occasional gibbon, and is typical of the larger indriids. Consequently, one can no longer accept any contention that the univertebral first costal capitular joint of A. afarensis implies that it did not use its upper limbs for locomotion. Rather, the formation of this joint is correlated with orthogrady and body size. We discuss a possible explanation of this correlation in terms of movement of the first rib during breathing in an orthograde primate, be it one that stands on two legs, swings by two arms, or clings to trunks using all four appendages.     

Subnasal alveolar morphology and the systematic position of Sivapithecus

Recent collecting in the Potwar Plateau of Pakistan has produced several new maxillae attributable to Sivapithecus. Since the subnasal region is preserved in most of these specimens, comparisons with early Miocene hominoid and Pliocene hominid maxillae become possible. On the basis of these comparisons, it has become clear that subnasal/premaxillary morphology distinguishes Asian and African hominoids. Ramapithecus and Sivapithecus share with Pongo an "Asian" subnasal pattern. The Proconsul species from the early Miocene of western Kenya and Australopithecus afarensis from the Hadar Formation of Ethiopia present two subsets of an "African" subnasal pattern. We think it likely that Ramapithecus and Sivapithecus represent a lineage that postdates the last common ancestor of African and Asian hominoids.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Evolution of the hominoid vertebral column: The long and the short of it

The postcranial axial skeleton exhibits considerable morphological and functional diversity among living primates. Particularly striking are the derived features in hominoids that distinguish them from most other primates and mammals. In contrast to the primitive catarrhine morphotype, which presumably possessed an external (protruding) tail and emphasized more pronograde trunk posture, all living hominoids are characterized by the absence of an external tail and adaptations to orthograde trunk posture. Moreover, modern humans evolved unique vertebral features that satisfy the demands of balancing an upright torso over the hind limbs during habitual terrestrial bipedalism. Our ability to identify the evolutionary timing and understand the functional and phylogenetic significance of these fundamental changes in postcranial axial skeletal anatomy in the hominoid fossil record is key to reconstructing ancestral hominoid patterns and retracing the evolutionary pathways that led to living apes and modern humans. Here, we provide an overview of what is known about evolution of the hominoid vertebral column, focusing on the currently available anatomical evidence of three major transitions: tail loss and adaptations to orthograde posture and bipedal locomotion.     

From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).     

Evaluation of Breathing Pattern: Comparison of a Manual Assessment of Respiratory Motion (MARM) and Respiratory Induction Plethysmography

Altered breathing pattern is an aspect of dysfunctional breathing but few standardised techniques exist to evaluate it. This study investigates a technique for evaluating and quantifying breathing pattern, called the Manual Assessment of Respiratory Motion (MARM) and compares it to measures performed with Respiratory Induction Plethysmography (RIP). About 12 subjects altered their breathing and posture while 2 examiners assessed their breathing using the MARM. Simultaneous measurements with RIP were taken. Inter-examiner agreement and agreement between MARM and RIP were assessed. The ability of the measurement methods to differentiate between diverse breathing and postural patterns was compared. High levels of agreement between examiners were found with the MARM for measures of the upper rib cage relative to lower rib cage/abdomen motion during breathing but not for measures of volume. The measures of upper rib cage dominance during breathing correlated with similar measures obtained from RIP. Both RIP and MARM measures methods were able to differentiate between abdominal and thoracic breathing patterns, but only MARM was able to differentiate between breathing changes occurring as result of slumped versus erect sitting posture. This study suggests that the MARM is a reliable clinical tool for assessing breathing pattern.     

New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus

A new complete hallucal metatarsal (SKX 5017) was recovered from the "lower bank" of Member 1 at Swartkrans (ca. 1.8 m.y. BP). The new metatarsal is attributed to Paranthropus robustus, the predominant hominid found in Member 1 (greater than 95% of hominid individuals). SKX 5017 is similar to Olduvai Hominid 8-H from bed I, Olduvai (ca. 1.76 m.y. BP), and both resemble humans most closely among extant hominoids. The base, shaft, and head of SKX 5017 suggest human-like foot posture and a human-like range of extension (= dorsiflexion) at the hallucal metatarsophalangeal joint, while at the same time the distal articular surface indicates that a human-like toe-off mechanism was absent in Paranthropus. The fossil evidence suggests that Homo habilis and Paranthropus may have attained a similar grade of bipedality at roughly 1.8 m.y. BP.     

Activity of respiratory muscles in upright and recumbent humans

It is established that during tidal breathing the rib cage expands more than the abdomen in the upright posture, whereas the reverse is usually true in the supine posture. To explore the reasons for this, we studied nine normal subjects in the supine, standing, and sitting postures, measuring thoracoabdominal movement with magnetometers and respiratory muscle activity via integrated electromyograms. In eight of the subjects, gastric and esophageal pressures and diaphragmatic electromyograms via esophageal electrodes were also measured. In the upright postures, there was generally more phasic and tonic activity in the scalene, sternocleidomastoid, and parasternal intercostal muscles. The diaphragm showed more phasic (but not more tonic) activity in the upright postures, and the abdominal oblique muscle showed more tonic (but not phasic) activity in the standing posture. Relative to the esophageal pressure change with inspiration, the inspiratory gastric pressure change was greater in the upright than in the supine posture. We conclude that the increased rib cage motion characteristic of the upright posture owes to a combination of increased activation of rib cage inspiratory muscles plus greater activation of the diaphragm that, together with a stiffened abdomen, acts to move the rib cage more effectively.     

The eruption pattern of the permanent incisors and first permanent molars in Australopithecus (Paranthropus) robustus

This study aims to reassess the claim that the eruption sequence of the permanent incisor and first permanent molar teeth of Australopithecus (Paranthropus) robustus is identical with that in modern Homo sapiens. Eight fossil hominid mandibles of equivalent dental developmental age were chosen for comparative study. Emphasis has been placed upon the comparative timing of events within the growth period rather than eruption sequence alone. The results of this study indicate that Homo sapiens and Australopithecus (Paranthropus) robustus share the same pattern of permanent molar and incisor eruption and that this is significantly different from the pattern of eruption shared by the great apes, Australopithecus africanus and Australopithecus afarensis.     

The morphology of the gluteus maximus during human evolution: Prerequisite or consequence of the upright bipedal posture?

The human gluteus maximus differs from that of the other hominoids because of its size and bony attachments. These differences raise questions concerning their sequence of appearance in human evolution. Given that humans practice a unique locomotor style, one wonders if the human gluteus maximus morphology is a prerequisite or a consequence of upright bipedal locomotion. This question is addressed using a computer model that evaluates muscle leverage in a variety of locomotor postures. In this model, the human-like, or ape-like, muscular pattern is imposed upon a representative hindlimb of each of the five extant hominoids. Shapes of the skeletal elements (i.e. ilium and ischium lengths) are adjusted in the computer to simulate an evolutionary progression from an ape to a human skeletal morphology. Changes in the leverage of different parts of the gluteus maximus (measured as moment arms) are monitored during this transition. The results show how the mechanical leverages of the gluteus maximus would have changed in a variety of hypothetical evolutionary sequences that describe an ape to human transition. Although the hominoid models exhibit minor differences in these simulations, they all show that the postural and locomotor functions of the gluteus maximus would become more difficult if musculoskeletal morphology changed to the human-like pattern before erect bipedal posture was adopted. Conversely, small adjustments in the ape-like musculoskeletal condition support an erect bipedal posture. These results suggest that a human like posture would have preceded the appearance of the human-like musculoskeletal morphology. Human gluteal morphology, therefore, is a consequence and not a prerequisite of the upright bipedal posture.     

Forelimb segment length proportions in extant hominoids and Australopithecus afarensis

Forelimb proportions have been used to infer locomotor adaptation in Australopithecus afarensis. However, little is known about proportions among individual forelimb segments in extant or fossil hominoids. The partial A. afarensis skeleton A.L. 438-1 and the more complete skeleton A.L. 288-1 provide the opportunity to assess relative length of the arm, forearm, wrist, and palm. We compare scaling relationships between pairs of forelimb bones of extant hominoids and A. afarensis, and length of individual forelimb elements to a body size surrogate. Hylobatids, and to a lesser extent orangutans, have the longest forelimb bones relative to size, although the carpus varies little among taxa, perhaps due to functional constraints of the wrist. Pan species are unique in having long metacarpals relative to ulnar length, demonstrating that they probably differ from the common chimp-human ancestor, and also that developmental mechanisms can be altered to results in differential growth of individual forelimb segments. A. afarensis has no forelimb bones that are significantly longer than those of humans for its size. It falls within the range of variation seen in modern humans for all comparisons relative to size, but appears to differ from the typical human brachial index due to a slightly shorter humerus and/or slightly longer ulna. It has short metacarpals like humans only among hominoids. Thus, while Pan may have elongated its metacarpus relative to ulnar length, A. afarensis may have reduced the length of its metacarpals and possibly its humerus relative to body size from the primitive condition.     

Hominid footprints at Laetoli: facts and interpretations

The history of discovery and interpretation of primate footprints at the site of Laetoli in northern Tanzania is reviewed. An analysis of the geological context of these tracks is provided. The hominid tracks in Tuff 7 at Site G in the Garusi River Valley demonstrate bipedality at a mid-Pliocene datum. Comparison of these tracks and the Hadar hominid foot fossils by Tuttle has led him to conclude that Australopithecus afarensis did not make the Tanzanian prints and that a more derived form of hominid is therefore indicated at Laetoli. An alternative interpretation has been offered by Stern and Susman who posit a conforming "transitional morphology" in both the Tanzanian prints and the Ethiopian bones. The present examines both hypotheses and shows that neither is likely to be entirely correct. To illustrate this point, a reconstruction of the foot skeleton of a female A. afarensis is undertaken, and the results are compared to the Laetoli tracks. We conclude that A. afarensis represents the best candidate for the maker of the Laetoli hominid trails.     

The metabolic costs of 'bent-hip, bent-knee' walking in humans

The costs of different modes of bipedalism are a key issue in reconstructing the likely gait of early human ancestors such as Australopithecus afarensis. Some workers, on the basis of morphological differences between the locomotor skeleton of A. afarensis and modern humans, have proposed that this hominid would have walked in a 'bent-hip, bent-knee' (BHBK) posture like that seen in the voluntary bipedalism of untrained chimpanzees. Computer modelling studies using inverse dynamics indicate that on the basis of segment proportions AL-288-1 should have been capable of mechanically effective upright walking, but in contrast predicted that BHBK walking would have been highly ineffective. The measure most pertinent to natural selection, however, is more likely to be the complete, physiological, or metabolic energy cost. We cannot measure this parameter in a fossil. This paper presents the most complete investigation yet of the metabolic and thermoregulatory costs of BHBK walking in humans. Data show that metabolic costs including the basal metabolic rate (BMR) increase by around 50% while the energy costs of locomotion and blood lactate production nearly double, heat load is increased, and core temperature does not return to normal within 20 minutes rest. Net effects imply that a resting period of 150% activity time would be necessary to prevent physiologically intolerable heat load. Preliminary data for children suggest that scaling effects would not significantly reduce relative costs for hominids of AL-288-1's size. Data from recent studies using forwards dynamic modelling confirm that similar total (including BMR) and locomotor metabolic costs would have applied to BHBK walking by AL-288-1. We explore some of the ecological consequences of our findings.     

On the systematic status of the late Neogene hominoids from Yunnan Province,China

Late Miocene and Pliocene hominoids from Yunnan Province in southern China have been recovered from four sites or site complexes: Xiaolongtan, Yangyi, Shihuiba and Yuanmou. Of these, Shihuiba and Yuanmou are among the most prolific fossil hominoid sites in Eurasia, and they have yielded important evidence that is critical for documenting the evolutionary history, biogeography and paleobiology of later Neogene hominids. The aim of this paper is to clarify their taxonomy and nomenclature, and to present a preliminary synthesis of their phylogenetic relationships and biogeography. The morphological pattern and degree of variation observed in the fossil samples is consistent with there being a single, sexually dimorphic species represented at each site. Provisionally, we consider the Shihuiba, Xiaolongtan and Yuanmou samples to belong to two separate species within a single genus. The valid names for these species are Lufengpithecus lufengensis (from Shihuiba) and L. keiyuanensis (from Xiaolongtan and Yuanmou). From a phylogenetic perspective, the currently available evidence suggests that Lufengpithecus is either a primitive hominid that represents the sister taxon of the Ponginae+Homininae or a primitive sister taxon to the Ponginae. We tend to favor the second alternative, but acknowledge that a more comprehensive comparative analysis is needed to substantiate the phylogenetic and taxonomic affinities of Lufengpithecus. Importantly, the Yunnan fossil apes provide a unique temporal perspective on the evolutionary history of hominoids. Their continued occurrence during the late Miocene and Pliocene (approximately 8-2Ma), when hominoids became extinct throughout the rest of Eurasia, suggests that southern China (and presumably southeast Asia in general) was an important refugium for hominoids, including the ancestors of the orang-utans and gibbons. The uplift of the Tibetan plateau and its impact on regional climatic conditions may have been an important contributing factor in isolating the hominoids geographically and ecologically. We speculate that changed climatic condition in the mid-Pliocene, and possibly the arrival of Homo soon after, may have precipitated the regional extinction of large hominoids in southern China and in mainland southeast Asia.     

Evolution of the power ("squeeze") grip and its morphological correlates in hominids.

The "squeeze" form of power grip is investigated for the purposes of clarifying the hand posture and activities associated with the grip, assessing the potential in chimpanzees for using the grip, and identifying morphological correlates of an effective power grip that may be recognized in fossil hominid species. Our approaches include: (1) the analysis of the human grip, focusing on both the hand posture involved and hand movements associated with use of the grip in hammering; (2) the analysis of similar chimpanzee grips and associated movements; (3) comparative functional analysis of regions in the hand exploited and stressed by the grip and its associated movements in humans; and (4) a review of the literature on the power grip and its morphological correlates. Results of the study indicate that humans use a squeeze form of power grip effectively to wield cylindrical tools forcefully as extensions of the forearm. Several morphological features occur in high frequency among humans which facilitate the grip and are consistent with the large internal and external forces associated with it in hammering and in other tool-using activities. Chimpanzee hand postures resembling this form of human power grip are not fully comparable and lack some of these morphological correlates that facilitate its use. The hand of Australopithecus afarensis does not appear to have been stressed by use of the grip, but there is some evidence for this type of stress in the metacarpals from Sterkfontein Member 4. Hands from Olduvai and Swartkrans do not provide sufficient evidence for assessment of power grip capabilities.     

Morphological study of the anthropoid thoracic cage: scaling of thoracic width and an analysis of rib curvature

While a relatively broad thorax and strongly curved ribs are widely regarded as common features of living hominoids, few studies have quantitatively examined these traits by methods other than calculating the chest index. The present study aims to quantify variations in thoracic cage morphology for living anthropoids. The odd-numbered ribs (first to eleventh) were articulated with the corresponding vertebrae and the cranial and lateral views subsequently photographed. Rib profiles were digitized in both views and line-fitted by a Bézier curve to create a three-dimensional morphological data set. When thoracic cage width was scaled against body mass, Hylobates (and possibly Pongo) plotted above non-hominoid anthropoids at almost all rib levels, while Pan did not differ from non-hominoid anthropoids. The overall pattern of the normalized thoracic width differed between Hylobates and other hominoids. In Hylobates, an upward convex curve was seen between the first and seventh ribs while a more linear pattern was observed in Pan and Pongo. This result quantitatively confirmed that the barrel-shaped thoracic cage in Hylobates can be distinguished from the funnel-shaped form in other hominoids. Conversely, all hominoids shared two distinct features in the upper half-thorax: (1) a pronounced dorsal protrusion of the proximal part of the rib in accordance with ventral displacement of the thoracic spine and (2) a relatively medially projecting sternal end. Although these features are likely to provide some mechanical advantage in orthograde and/or suspensory positional behaviors, they were barely present in the suspensory Ateles. An erratum to this article can be found at     

Species,speciation and human evolution

Human evolution is considered from the perspective of the recognition concept of species, which views species as an epiphenomenon of shared fertilisation systems in sexually reproducing organisms. It is argued that this concept predicts the controversial pattern of punctuated equilibrium, and offers an understanding of the hominid fossil evidence in line with that pattern. Changes in the nature of the fertilization system in the human lineage over time are discussed in relation to the pattern of morphological continuity between proposed species.     

Longgupo: EarlyHomo colonizer or late plioceneLufengpithecus survivor in south China?

A fragment of mandible and a maxillary incisor of different individuals from the Longgupo Cave, China have been cited as evidence of an early dispersal ofHomo from Africa to Asia. More specifically, these specimens are said to resemble “Homo ergaster” orHomo habilis, rather than the species usually thought to be the first Asian colonizer,Homo erectus. If this supposition is correct, it calls into question which hominid (sensu stricto) first left Africa, and why hominids became a colonizing species. Furthermore, the Longgupo remains have been used to buttress the argument thatHomo erectus evolved uniquely in Asia and was not involved in the origins of modern humans. We question this whole line of argument because the mandibular fragment cannot be distinguished from penecontemporary fossil apes, especially the Late Miocene-Pliocene Chinese genusLufengpithecus, while the incisor is indistinguishable from those of recent and living east Asian people and may be intrusive in the deposit. We believe that the Longgupo mandible represents the relic survival of a Late Miocene ape lineage into a period just prior to the dispersal of hominids into southeastern Asia, with some female dental features that parallel the hominid condition. If the Longgupo mandibular fragment represents a member of theLufengpithecus clade, it demonstrates that hominoids other thanGigantopithecus and the direct ancestor of the orangutan persisted in east Asia into the Late Pliocene, while all other Eurasian large-bodied hominoids disappeared in the Late Miocene.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。