Early Hominin Fossil Research Articles

Measures of maturation in early fossil hominins: events at the first transition from australopiths to early Homo.

An important question in palaeoanthropology is whether, among the australopiths and the first fossil hominins attributed to early Homo, there was a shift towards a more prolonged period of growth that can be distinguished from that of the living great apes and whether between the end of weaning and the beginning of puberty there was a slow period of growth as there is in modern humans. Evidence for the pace of growth in early fossil hominins comes from preserved tooth microstructure. A record of incremental growth in enamel and dentine persists, which allows us to reconstruct tooth growth and compare key measures of dental maturation with modern humans and living great apes. Despite their diverse diets and way of life, it is currently difficult to identify any clear differences in the timing of dental development among living great apes, australopiths and the earliest hominins attributed to the genus Homo There is, however, limited evidence that some early hominins may have attained a greater proportion of their body mass and stature relatively earlier in the growth period than is typical of modern humans today.This article is part of the themed issue 'Major transitions in human evolution'.

Scapular shape of extant hominoids and the African ape/modern human last common ancestor

Newly discovered early hominin fossil scapulae have bolstered investigations of scapular shape, which have long been used to interpret behavioral variation among primates. However, unexpected similarities between Pongo and Homo – particularly in scapular spine orientation – have raised questions about the functional utility of scapular morphology and its phylogenetic context in the hominin lineage. Not surprisingly, significant disagreement surrounds disparate morphological reconstructions of the modern human/African ape last common ancestor (LCA).Our study utilizes geometric morphometric (GM) approaches – two employing homologous, anatomical landmarks and a “spine-free” alternative using 98 sliding semilandmarks along the boundary of the subscapular fossa. The landmark-based “wireframe” GM analysis principally sorted groups by spine orientation: Homo and Pongo were similar to one another with more transversely-oriented spines as compared to Hylobates and the African apes. In contrast, Homo and Gorilla clustered together in our semilandmark analysis with superoinferiorly broad blades. Pan scapulae were similar, but had more mediolaterally compressed blades and laterally-positioned superior angles. Hylobates was superoinferiorly narrow, yet obliquely expanded relative to the vertebral border. Pongo scapulae were unique among hominoids in being nearly as broad as they were long.Previously documented ‘convergence’ of Homo and Pongo scapulae appears to be principally driven by similarities in spine orientation, rather than overall blade shape. Therefore, we contend that it is more parsimonious to reconstruct the African ape/Homo LCA scapula as being Gorilla-like, especially in light of similar characterizations of certain fossil hominin scapulae. Accordingly, the evolution of Pan (highly oblique spine and laterally-situated superior angle) and Homo (transversely-oriented spine) scapular morphology would have involved relatively minor shifts from this ancestral condition. These results support the prevailing molecular phylogeny and provide further insight into the behavioral implications of scapular shape in the LCA and fossil hominins.

Vertebral numbers and human evolution.

Ever since Tyson (1699), anatomists have noted and compared differences in the regional numbers of vertebrae among humans and other hominoids. Subsequent workers interpreted these differences in phylogenetic, functional, and behavioral frameworks and speculated on the history of vertebral numbers during human evolution. Even in a modern phylogenetic framework and with greatly expanded sample sizes of hominoid species, researchers' conclusions vary drastically, positing that hominins evolved from either a "long-backed" (numerically long lumbar column) or a "short-backed" (numerically short lumbar column) ancestor. We show that these disparate interpretations are due in part to the use of different criteria for what defines a lumbar vertebra, but argue that, regardless of which lumbar definition is used, hominins are similar to their great ape relatives in possessing a short trunk, a rare occurrence in mammals and one that defines the clade Hominoidea. Furthermore, we address the recent claim that the early hominin thoracolumbar configuration is not distinct from that of modern humans and conclude that early hominins show evidence of "cranial shifting," which might explain the anomalous morphology of several early hominin fossils. Finally, we evaluate the competing hypotheses on numbers of vertebrae and argue that the current data support a hominin ancestor with an African ape-like short trunk and lower back.

Age estimation in fossil hominins: comparing dental development in early Homo with modern humans

Background: Previous studies have used molar tooth eruption as a comparative marker of maturation in early fossil hominins. However, tooth eruption and tooth formation are independent maturational processes.Aim: To determine whether estimates of age for entering a stage of dental development in three early hominin fossils fell within the distribution of a modern human sample.Methods: This study used a comparative model of dental development to identify the stages of dental development most likely to provide information about length of the growth period in early fossil hominins. Age estimates for stages of dental development in fossils were superimposed onto a normal distribution of the same radiographically defined stages derived from a sample of 6540 children of diverse geographical origin.Results: Both within the dentition of S7–37, from Sangiran, Java, but also for stages of two other specimens (KNM-WT 15000 from Kenya and StW 151 from South Africa), all age estimates for later stages of tooth formation fell within the modern sample range.Conclusions: A pattern appears to exist in early Homo where, both within and between developing dentitions, age estimates for stages of P4, M2 and M3 tooth formation fell consistently among the more advanced individuals of the modern human sample.

Recent origin of low trabecular bone density in modern humans

Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

Middle Pliocene hominin mandibular fourth premolars from Woranso-Mille (Central Afar, Ethiopia)

The Woranso-Mille study area has thus far yielded more than 120 early hominin fossil specimens dated to between 3.4 and 3.8 million years ago. Previous studies indicate that dentognathic fossil remains from the study area show a mosaic of features shared by both Australopithecus anamensis and Australopithecus afarensis. Here, we describe 12 isolated mandibular fourth premolars recovered from the Woranso-Mille study area and compare them with those of other early hominins using both traditional comparative methods and geometric morphometric methods. The results indicate that the Woranso-Mille sample is most similar to Au. afarensis among hominins. However, some specimens show distinctive features of crown shape, namely an extremely bulging distolingual corner associated with a relatively large talonid and a mesiodistally elongated crown. This unique morphology is accompanied by a root form that is different from those identified previously. The existence of distinctive P4 morphology is intriguing given the presence of more than one hominin species at Woranso-Mille, although support for taxonomic heterogeneity in this sample is equivocal. Further, the taxonomic significance of these features is unclear, as they fail to distinguish Pan from Gorilla and known hominin species from one another.

Ecomorphology of radii in Canidae (Carnivora): application to fragmentary fossils from Plio-Pleistocene hominin assemblages

Fragmentary long bone material from fossil Carnivora is rarely considered to support palaeoenvironmental reconstructions. Here, we use morphometry of the radius in extant carnivorans of the dog family (Canidae) to reconstruct the palaeobiology of extinct canids from Olduvai Gorge, Tanzania (Bed I and II) and Koobi Fora, Kenya. We use radius morphometrics to predict adaptation to prey size and introduce a new method for quantifying canid habitat adaptations based on the geographic distributions of the extant species sampled. Linear Discriminant Function Analyses (DFA) and cluster neighbour-joining algorithms are employed to investigate radial morphometrics as described by 29 linear measurements. Results of our analyses suggest that a phylogenetic signal is present in radial morphometrics, even if it does not allow us to accurately discriminate among genera. A binary prey size categorisation of “small-medium” versus “large” prey can be more accurately predicted than a habitat categorisation scheme (Open, Mixed, Closed). The East African fossil specimens examined show morphometric affinities with the golden jackal (Canis aureus) and coyote (Canis latrans) and are likely attributable to the genus Canis. Fragmentary fossil specimens from Olduvai Gorge are predicted as habitat generalists (Open for Bed I and Mixed for Bed II) adapted for hunting small-medium prey, whereas the specimen from Koobi Fora was predicted as inhabiting mixed habitats and adapted for killing large prey. This study supports the inclusion of fossil Canidae in palaeoecological analyses attempting to clarify the palaeoenvironment of early hominin fossil sites.

The Hairless Mutation Hypothesis Explains Not Only the Origin of Humanization from the Human/Ape Common Ancestor but also Immature Baby Delivery

Molecular and paleontological dating indicates human appearance 6 million years ago. Early hominin fossils show that they were bipeds. Three salient characteristics distinguish humans from other primates: bipedality, practical nakedness, and the family reproductive unit. Once a hairless mutation was initially introduced, the three characteristics became separately inexplicable. All primates except humans can carry their babies without using their hands. A hairless mother would be forced to stand and walk upright to hold a baby. Her activities would be markedly limited. The male partner would have to collect food and carry it to her to keep their baby from starving; irresponsible and selfish males could not have left their offspring. The mother would have sexually accepted her partner at any time as a reward for food. Sexual relations irrespective of estrus cycles might have strengthened the pair bond. Consequently, hairless and upright pairs would have established strong families. Early hominins had the opposable hallux and remained as arboreal denizens. Climate changes probably forced them to terrestrial life, but the ground was full of danger and trees were indispensable for refuge and nesting. Consequently, archaic hominins had mosaic characteristics of the upper body adapted for arboreal life and the lower body for terrestrial life, for which a larger brain became advantageous. Alternative strategies became possible: development of a large pelvis with a big birth canal through which a baby with a big head could pass, or delivery of an immature baby, with rearing after birth. The former was physically incompatible with an upright posture, and structurally unfavorable for swift movement. The latter was unavailable to primates, the babies of which had to cling to the mother; upright hominins were able to hold the immature baby with hands and raise it after birth.

Exploring Third Metacarpal Capitate Facet Shape in Early Hominins

The joint between the capitate and third metacarpal plays an important role in stabilizing the manus during hand use in great apes and humans. Researchers have examined the morphology of this region in humans, our fossil relatives, and other extant primates to try to understand the importance of this joint in human evolution. The first goal of our research was to explore shape variation of the third metacarpal capitate facet across extant anthropoids, including hominoids, cercopithecoids, and platyrrhines. This analysis allowed us to examine the range of variation in the capitate facet and the degree to which locomotor behavior, phylogeny, and size explained shape variation. We also examined capitate facet shape in the early hominin fossil record in order to explore how the shape of this articular surface has changed during early hominin evolution. We captured six landmark coordinates on the edge of the capitate facet in extant anthropoids and fossil specimens to quantify and visualize shape variation in this region. We used principal components analysis, Procrustes distances, and multivariate regression analysis to investigate different possible influences on shape variation. We found that shape variation corresponded to function, phylogeny, and size. With the exception of brachiation, shape variation did not clearly correspond with any specific locomotor behavior. However, we identified a shift in the relative mediolateral breadth of the capitate facet during early hominin evolution, which is most likely one of several adaptations for a more stable joint surface.

Hairless mutation: a driving force of humanization from a human–ape common ancestor by enforcing upright walking while holding a baby with both hands

Three major characteristics distinguish humans from other primates: bipedality, practical nakedness, and the family as a social unit. A hairless mutation introduced into the chimpanzee/human last common ancestor (CLCA) 6 million years ago (Mya) diverged hairless human and hairy chimpanzee lineages. All primates except humans can carry their babies without using their hands. A hairless mother would be forced to stand and walk upright. Her activities would be markedly limited. The male partner would have to collect food and carry it to her by hand to keep her and their baby from starving; irresponsible and selfish males could not have left their offspring. The mother would have sexually accepted her partner at any time as a reward for food. Sexual relations irrespective of estrus cycles might have strengthened the pair bond. Molecular and paleontological dating indicates that CLCA existed 6 Mya, and early hominin fossils show that they were bipeds, indicating that humanization from CLCA occurred rapidly. A single mutation in animals with scalp hair is known to induce hairless phenotype (ectodermal dysplasia). Bipedalism and hairlessness are disadvantageous traits; only those who could survive trials and tribulations in cooperation with family members must have been able to evolve as humans.

The Omo‐Turkana Basin Fossil Hominins and Their Contribution to Our Understanding of Human Evolution in Africa

The Omo-Turkana Basin, including the hominin fossil sites around Lake Turkana and the sites along the lower reaches of the Omo River, has made and continues to make an important contribution to improving our murky understanding of human evolution. This review highlights the various ways the Omo-Turkana Basin fossil record has contributed to, and continues to challenge, interpretations of human evolution. Despite many diagrams that look suspiciously like comprehensive hypotheses about human evolutionary history, any sensible paleoanthropologist knows that the early hominin fossil record is too meager to do anything other than offer very provisional statements about hominin taxonomy and phylogeny. If history tells us anything, it is that we still have much to learn about the hominin clade. Thus, we summarize the current state of knowledge of the hominin species represented at the Omo-Turkana Basin sites. We then focus on three specific topics for which the fossil evidence is especially relevant: the origin and nature of Paranthropus; the origin and nature of early Homo; and the ongoing debate about whether the pattern of human evolution is more consistent with speciation by cladogenesis, with greater taxonomic diversity or with speciation by anagenetic transformation, resulting in less taxonomic diversity and a more linear interpretation of human evolutionary history.

Retrieving chronological age from dental remains of early fossil hominins to reconstruct human growth in the past.

A chronology of dental development in Pan troglodytes is arguably the best available model with which to compare and contrast reconstructed dental chronologies of the earliest fossil hominins. Establishing a time scale for growth is a requirement for being able to make further comparative observations about timing and rate during both dento-skeletal growth and brain growth. The absolute timing of anterior tooth crown and root formation appears not to reflect the period of somatic growth. In contrast, the molar dentition best reflects changes to the total growth period. Earlier initiation of molar mineralization, shorter crown formation times, less root length formed at gingival emergence into functional occlusion are cumulatively expressed as earlier ages at molar eruption. Things that are similar in modern humans and Pan, such as the total length of time taken to form individual teeth, raise expectations that these would also have been the same in fossil hominins. The best evidence there is from the youngest fossil hominin specimens suggests a close resemblance to the model for Pan but also hints that Gorilla may be a better developmental model for some. A mosaic of great ape-like features currently best describes the timing of early hominin dental development.

New magnetostratigraphy for the Olduvai Subchron in the Koobi Fora Formation, northwest Kenya, with implications for early Homo

A problematic magnetostratigraphy for the Koobi Fora Formation has contributed to debates on the evolutionary implications for early hominin fossils. To address this, 50 independent samples distributed over a nearly 63-m-thick interval were collected from the lower-middle KBS Member type section in fossil collection Area 102, northeast Turkana Basin. Characteristic directions obtained by thermal demagnetization define a coherent magnetostratigraphy that is supported by alternating-field studies on 28 sister specimens and the prior tephrochronological framework. Two long polarity intervals were recognized, each 30–40 m in thickness, and interpreted as the upper part of the normal polarity Olduvai Subchron and the overlying reverse polarity Matuyama Chron. The end Olduvai consists of a normal–reverse–normal polarity sequence occurring over a thickness of at least 1 m but perhaps up to 5 m, suggesting that this subchron has a short reverse interval in its uppermost part. Such a fine-scale structure also has been reported from several other sites, like the Pliocene–Pleistocene boundary and point stratotype section at Vrica, Italy, which serves as a basis for formally delimiting three temporally discrete polarity subintervals for the Olduvai Subchron. These paleomagnetic results that place the upper boundary of the Olduvai at ∼ 48 m above the base of the KBS Member, coupled with published radioisotopic dates, firmly secure the age of partial cranium KNM-ER 3733 in the interval 1.78–1.48 Ma, with an interpolated age of ∼ 1.7 Ma, giving this fossil the most unambiguous numerical-age constraints, as compared to the oldest Homo cranial remains from Europe and Asia. Nonetheless, assured placement of the top of the Olduvai Subchron in the KBS Member is not sufficient in the face of other uncertainties to influence conventional interpretations of the timing and direction for the global dispersal of early Homo erectus.

Functional morphology of the ankle and the likelihood of climbing in early hominins

Whether early hominins were adept tree climbers is unclear. Although some researchers have argued that bipedality maladapts the hominin skeleton for climbing, others have argued that early hominin fossils display an amalgamation of features consistent with both locomotor strategies. Although chimpanzees have featured prominently in these arguments, there are no published data on the kinematics of climbing in wild chimpanzees. Without these biomechanical data describing how chimpanzees actually climb trees, identifying correlates of climbing in modern ape skeletons is difficult, thereby limiting accurate interpretations of the hominin fossil record. Here, the first kinematic data on vertical climbing in wild chimpanzees are presented. These data are used to identify skeletal correlates of climbing in the ankle joint of the African apes to more accurately interpret hominin distal tibiae and tali. This study finds that chimpanzees engage in an extraordinary range of foot dorsiflexion and inversion during vertical climbing bouts. Two skeletal correlates of modern ape-like vertical climbing are identified in the ankle joint and related to positions of dorsiflexion and foot inversion. A study of the 14 distal tibiae and 15 tali identified and published as hominins from 4.12 to 1.53 million years ago finds that the ankles of early hominins were poorly adapted for modern ape-like vertical climbing bouts. This study concludes that if hominins included tree climbing as part of their locomotor repertoire, then they were performing this activity in a manner decidedly unlike modern chimpanzees.

Brief communication: Enamel thickness trends in the dental arcade of humans and chimpanzees

In addition to evidence for bipedality in some fossil taxa, molar enamel thickness is among the few characters distinguishing (thick-enameled) hominins from the (thin-enameled) African apes. Despite the importance of enamel thickness in taxonomic discussions and a long history of scholarship, measurements of enamel thickness are performed almost exclusively on molars, with relatively few studies examining premolars and anterior teeth. This focus on molars has limited the scope of enamel thickness studies (i.e., there exist many fossil hominin incisors, canines, and premolars). Increasing the available sample of teeth from which to compare enamel thickness measurements from the fossil record could substantially increase our understanding of this aspect of dental biology, and perhaps facilitate greater taxonomic resolution of early hominin fossils. In this study, we report absolute and relative (size-scaled) enamel thickness measurements for the complete dentition of modern humans and chimpanzees. In accord with previous studies of molars, chimpanzees show lower relative enamel thickness at each tooth position, with little overlap between the two taxa. A significant trend of increasing enamel thickness from anterior to posterior teeth is apparent in both humans and chimpanzees, indicating that inter-taxon comparisons should be limited to the same tooth position in order to compare homologous structures. As nondestructive imaging techniques become commonplace (facilitating the examination of increasing numbers of fossil specimens), studies may maximize available samples by expanding beyond molars.

Chimpanzee locomotor energetics and the origin of human bipedalism

Bipedal walking is evident in the earliest hominins [Zollikofer CPE, Ponce de Leon MS, Lieberman DE, Guy F, Pilbeam D, et al. (2005) Nature 434:755-759], but why our unique two-legged gait evolved remains unknown. Here, we analyze walking energetics and biomechanics for adult chimpanzees and humans to investigate the long-standing hypothesis that bipedalism reduced the energy cost of walking compared with our ape-like ancestors [Rodman PS, McHenry HM (1980) Am J Phys Anthropol 52:103-106]. Consistent with previous work on juvenile chimpanzees [Taylor CR, Rowntree VJ (1973) Science 179:186-187], we find that bipedal and quadrupedal walking costs are not significantly different in our sample of adult chimpanzees. However, a more detailed analysis reveals significant differences in bipedal and quadrupedal cost in most individuals, which are masked when subjects are examined as a group. Furthermore, human walking is approximately 75% less costly than both quadrupedal and bipedal walking in chimpanzees. Variation in cost between bipedal and quadrupedal walking, as well as between chimpanzees and humans, is well explained by biomechanical differences in anatomy and gait, with the decreased cost of human walking attributable to our more extended hip and a longer hindlimb. Analyses of these features in early fossil hominins, coupled with analyses of bipedal walking in chimpanzees, indicate that bipedalism in early, ape-like hominins could indeed have been less costly than quadrupedal knucklewalking.

Earliest evidence of modern human life history in North African early Homo sapiens.

Recent developmental studies demonstrate that early fossil hominins possessed shorter growth periods than living humans, implying disparate life histories. Analyses of incremental features in teeth provide an accurate means of assessing the age at death of developing dentitions, facilitating direct comparisons with fossil and modern humans. It is currently unknown when and where the prolonged modern human developmental condition originated. Here, an application of x-ray synchrotron microtomography reveals that an early Homo sapiens juvenile from Morocco dated at 160,000 years before present displays an equivalent degree of tooth development to modern European children at the same age. Crown formation times in the juvenile's macrodont dentition are higher than modern human mean values, whereas root development is accelerated relative to modern humans but is less than living apes and some fossil hominins. The juvenile from Jebel Irhoud is currently the oldest-known member of Homo with a developmental pattern (degree of eruption, developmental stage, and crown formation time) that is more similar to modern H. sapiens than to earlier members of Homo. This study also underscores the continuing importance of North Africa for understanding the origins of human anatomical and behavioral modernity. Corresponding biological and cultural changes may have appeared relatively late in the course of human evolution.

Revised age estimates of Australopithecus-bearing deposits at Sterkfontein, South Africa.

The Sterkfontein fossil site in South Africa has produced the largest concentration of early hominin fossils from a single locality. Recent reports suggest that Australopithecus from this site is found within a broad paleontological age of between 2.5-3.5 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130; Kuman and Clarke [2000] J Hum Evol 38:827-847). Specifically, the hominin fossil commonly referred to as the "Little Foot" skeleton from Member 2, which is arguably the most complete early hominin skeleton yet discovered, has been magnetostratigraphically dated to 3.30-3.33 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130). More recent claims suggest that hominin fossils from the Jacovec Cavern are even older, being dated to approximately 3.5 Ma. Our interpretation of the fauna, the archeometric results, and the magnetostratigraphy of Sterkfontein indicate that it is unlikely that any Members yet described from Sterkfontein are in excess of 3.04 Ma in age. We estimate that Member 2, including the Little Foot skeleton, is younger than 3.0 Ma, and that Member 4, previously dated to between 2.4-2.8 Ma, is more likely to fall between 1.5-2.5 Ma. Our results suggest that Australopithecus africanus should not be considered as a temporal contemporary of Australopithecus afarensis, Australopithecus bahrelghazali, and Kenyanthropus platyops.