Footprint evidence of early hominin locomotor diversity at Laetoli, Tanzania.

Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage1-3. Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity3-5. In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli.

A review of the distal femur in Australopithecus.

In 1938, the first distal femur of a fossil Australopithecus was discovered at Sterkfontein, South Africa. A decade later, another distal femur was discovered at the same locality. These two fossil femora were the subject of a foundational paper authored by Kingsbury Heiple and Owen Lovejoy in 1971. In this paper, the authors discussed functionally relevant anatomies of these two fossil femora and noted their strong affinity to the modern human condition. Here, we update this work by including eight more fossil Australopithecus distal femora, an expanded comparative dataset, as well as additional linear measurements. Just as Heiple and Lovejoy reported a half-century ago, we find strong overlap between modern humans and cercopithecoids, except for inferiorly flattened condyles and a high bicondylar angle, both of which characterize modern humans and Australopithecus and are directly related to striding bipedalism. All other measured aspects of the femora are by-products of these key morphological traits. Additional fossil material from the early Pliocene will help to inform the evolution of the hominin distal femur and its condition in the Pan-Homo common ancestor that preceded bipedal locomotion.

A new early hominin calcaneus from Kromdraai (South Africa).

The Kromdraai site in South Africa has yielded numerous early hominin fossils since 1938. As a part of recent excavations within Unit P, a largely complete early hominin calcaneus (KW 6302) was discovered. Due to its role in locomotion, the calcaneus has the potential to reveal important form/function relationships. Here, we describe KW 6302 and analyze its preserved morphology relative to human and nonhuman ape calcanei, as well as calcanei attributed to Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Homo naledi, and the Omo calcaneus (either Paranthropus or early Homo). KW 6302 calcaneal morphology is assessed using numerous quantitative metrics including linear measures, calcaneal robusticity index, relative lateral plantar process position, Achilles tendon length reconstruction, and a three-dimensional geometric morphometric sliding semilandmark analysis. KW 6302 exhibits an overall calcaneal morphology that is intermediate between humans and nonhuman apes, although closer to modern humans. KW 6302 possesses many traits that indicate it was likely well-adapted for terrestrial bipedal locomotion, including a relatively flat posterior talar facet and a large lateral plantar process that is similarly positioned to modern humans. It also retains traits that indicate that climbing may have remained a part of its locomotor repertoire, such as a relatively gracile tuber and a large peroneal trochlea. Specimens from Kromdraai have been attributed to either Paranthropus robustus or early Homo; however, there are no definitively attributed calcanei for either genus, making it difficult to taxonomically assign this specimen. KW 6302 and the Omo calcaneus, however, fall outside the range of expected variation for an extant genus, indicating that if the Omo calcaneus was Paranthropus, then KW 6302 would likely be attributed to early Homo (or vice versa).

Femoral neck and shaft structure in Homo naledi from the Dinaledi Chamber (Rising Star System, South Africa).

The abundant femoral assemblage of Homo naledi found in the Dinaledi Chamber provides a unique opportunity to test hypotheses regarding the taxonomy, locomotion, and loading patterns of this species. Here we describe neck and shaft cross-sectional structure of all the femoral fossils recovered in the Dinaledi Chamber and compare them to a broad sample of fossil hominins, recent humans, and extant apes. Cross-sectional geometric (CSG) properties from the femoral neck (base of neck and midneck) and diaphysis (subtrochanteric region and midshaft) were obtained through CT scans for H. naledi and through CT scans or from the literature for the comparative sample. The comparison of CSG properties of H. naledi and the comparative samples shows that H. naledi femoral neck is quite derived with low superoinferior cortical thickness ratio and high relative cortical area. The neck appears superoinferiorly elongated because of two bony pilasters on its superior surface. Homo naledi femoral shaft shows a relatively thick cortex compared to the other hominins. The subtrochanteric region of the diaphysis is mediolaterally elongated resembling early hominins while the midshaft is anteroposteriorly elongated, indicating high mobility levels. In term of diaphyseal robusticity, the H. naledi femur is more gracile that other hominins and most apes. Homo naledi shows a unique combination of characteristics in its femur that undoubtedly indicate a species committed to terrestrial bipedalism but with a unique loading pattern of the femur possibly consequence of the unique postcranial anatomy of the species.

Morphology of the Homo naledi femora from Lesedi.

OBJECTIVES: The femoral remains recovered from the Lesedi Chamber are among the most complete South African fossil hominin femora discovered to date and offer new and valuable insights into the anatomy and variation of the bone in Homo naledi. While the femur is one of the best represented postcranial elements in the H. naledi assemblage from the Dinaledi Chamber, the fragmentary and commingled nature of the Dinaledi femoral remains has impeded the assessment of this element in its complete state. MATERIALS AND METHODS: Here we analyze and provide descriptions of three new relatively well-preserved femoral specimens of H. naledi from the Lesedi Chamber: U.W. 102a-001, U.W. 102a-003, and U.W. 102a-004. These femora are quantitatively and qualitatively compared to multiple extinct hominin femoral specimens, extant hominid taxa, and, where possible, each other. RESULTS: The Lesedi femora are morphologically similar to the Dinaledi femora for all overlapping regions, with differences limited to few traits of presently unknown significance. The Lesedi distal femur and mid-diaphysis preserve anatomy previously unidentified or unconfirmed in the species, including an anteroposteriorly expanded midshaft and anteriorly expanded patellar surface. The hypothesis that the Lesedi femoral sample may represent two individuals is supported. DISCUSSION: The Lesedi femora increase the range of variation of femoral morphology in H. naledi. Newly described features of the diaphysis and distal femur are either taxonomically uninformative or Homo-like. Overall, these three new femora are consistent with previous functional interpretations of the H. naledi lower limb as belonging to a species adapted for long distance walking and, possibly, running.

A quantification of calcaneal lateral plantar process position with implications for bipedal locomotion in Australopithecus.

The evolution of bipedalism in the hominin lineage has shaped the posterior human calcaneus into a large, robust structure considered to be adaptive for dissipating peak compressive forces and energy during heel-strike. A unique anatomy thought to contribute to the human calcaneus and its function is the lateral plantar process (LPP). While it has long been known that humans possess a plantarly positioned LPP and apes possess a more dorsally positioned homologous structure, the relative position of the LPP and intraspecific variation of this structure have never been quantified. Here, we present a method for quantifying relative LPP position and find that, while variable, humans have a significantly more plantar position of the LPP than that found in the apes. Among extinct hominins, while the position of the LPP in Australopithecus afarensis falls within the human distribution, the LPP is more dorsally positioned in Australopithecus sediba and barely within the modern human range of variation. Results from a resampling procedure suggest that these differences can reflect either individual variation of a foot structure/function largely shared among Australopithecus species, or functionally distinct morphologies that reflect locomotor diversity in Plio-Pleistocene hominins. An implication of the latter possibility is that calcaneal changes adaptive for heel-striking bipedalism may have evolved independently in two different hominin lineages.

The evolution of the human foot.

There are 26 bones in each foot (52 in total), meaning that roughly a quarter of the human skeleton consists of foot bones. Yet, early hominin foot fossils are frustratingly rare, making it quite difficult to reconstruct the evolutionary history of the human foot. Despite the continued paucity of hominid or hominin foot fossils from the late Miocene and early Pliocene, the last decade has witnessed the discovery of an extraordinary number of early hominin foot bones, inviting a reassessment of how the human foot evolved, and providing fresh new evidence for locomotor diversity throughout hominin evolution. Here, we provide a review of our current understanding of the evolutionary history of the hominin foot.

The thigh and leg of Homo naledi.

This paper describes the 108 femoral, patellar, tibial, and fibular elements of a new species of Homo (Homo naledi) discovered in the Dinaledi chamber of the Rising Star cave system in South Africa. Homo naledi possesses a mosaic of primitive, derived, and unique traits functionally indicative of a bipedal hominin adapted for long distance walking and possibly running. Traits shared with australopiths include an anteroposteriorly compressed femoral neck, a mediolaterally compressed tibia, and a relatively circular fibular neck. Traits shared with Homo include a well-marked linea aspera, anteroposteriorly thick patellae, relatively long tibiae, and gracile fibulae with laterally oriented lateral malleoli. Unique features include the presence of two pillars on the superior aspect of the femoral neck and a tubercular distal insertion of the pes anserinus on the tibia. The mosaic morphology of the H. naledi thigh and leg appears most consistent with a species intermediate between Australopithecus spp. and Homo erectus and, accordingly, may offer insight into the nature of the earliest members of genus Homo. These fossils also expand the morphological diversity of the Homo lower limb, perhaps indicative of locomotor diversity in our genus.

Virtual reconstruction of the Australopithecus africanus pelvis Sts 65 with implications for obstetrics and locomotion.

Characterizing australopith pelvic morphology has been difficult in part because of limited fossilized pelvic material. Here, we reassess the morphology of an under-studied adult right ilium and pubis (Sts 65) from Member 4 of Sterkfontein, South Africa, and provide a hypothetical digital reconstruction of its overall pelvic morphology. The small size of the pelvis, presence of a preauricular sulcus, and shape of the sciatic notch allow us to agree with past interpretations that Sts 65 likely belonged to a female. The morphology of the iliac pillar, while not as substantial as in Homo, is more robust than in A.L. 288-1 and Sts 14. We created a reconstruction of the pelvis by digitally articulating the Sts 65 right ilium and a mirrored copy of the left ilium with the Sts 14 sacrum in Autodesk Maya. Points along the arcuate line were used to orient the ilia to the sacrum. This reconstruction of the Sts 65 pelvis looks much like a "classic" australopith pelvis, with laterally flared ilia and an inferiorly deflected pubis. An analysis of the obstetric dimensions from our reconstruction shows similarity to other australopiths, a likely transverse or oblique entrance of the neonatal cranium into the pelvic inlet, and a cephalopelvic ratio similar to that found in humans today.

The relationship between foot arch measurements and walking parameters in children.

BACKGROUND: Walking mechanics are influenced by body morphology. Foot arch height is one aspect of body morphology central to walking. However, generalizations about the relationship between arch height and walking are limited due to previous methodologies used for measuring the arch and the populations that have been studied. To gain the knowledge needed to support healthy gait in children and adults, we need to understand this relationship in unimpaired, typically developing children and adults using dynamic measures. The purpose of the current study was to examine the relationship between arch height and gait in a sample of healthy children and adults using dynamic measures. METHODS: Data were collected from 638 participants (n = 254 children and n = 384 adults) at the Museum of Science, Boston (MOS) and from 18 4- to 8-year-olds at the Motor Development and Motor Control Laboratories. Digital footprints were used to calculate two arch indices: the Chippaux-Smirak (CSI) and the Keimig Indices (KI). The height of the navicular bone was measured. Gait parameters were captured with a mechanized gait carpet at the MOS and three-dimensional motion analyses and in-ground force plates in the Motor Development and Motor Control Laboratories. RESULTS: Linear regression analyses on data from the MOS confirmed that as age increases, step length increases. With a linear mixed effect regression model, we found that individuals who took longer steps had higher arches as measured by the KI. However, this relationship was no longer significant when only adults were included in the model. A model restricted to children found that amongst this sample, those with higher CSI and higher KI values take longer relative step lengths. Data from the Motor Development and Motor Control Laboratories showed that both CSI and KI added to the prediction; children with lower anterior ground reaction forces had higher CSI and higher KI values. Arch height indices were correlated with navicular height. CONCLUSIONS: These results suggest that more than one measure of the arch may be needed elucidate the relationship between arch height and gait.

A neonatal perspective on Homo erectus brain growth.

The Mojokerto calvaria has been central to assessment of brain growth in Homo erectus, but different analytical approaches and uncertainty in the specimen's age at death have hindered consensus on the nature of H. erectus brain growth. We simulate average annual rates (AR) of absolute endocranial volume (ECV) growth and proportional size change (PSC) in H. erectus, utilizing estimates of H. erectus neonatal ECV and a range of ages for Mojokerto. These values are compared with resampled ARs and PSCs from ontogenetic series of humans, chimpanzees, and gorillas from birth to six years. Results are consistent with other studies of ECV growth in extant taxa. There is extensive overlap in PSC between all living species through the first postnatal year, with continued but lesser overlap between humans and chimpanzees to age six. Human ARs are elevated above those of apes, although there is modest overlap up to 0.50 years. Ape ARs overlap throughout the sequence, with gorillas slightly elevated over chimpanzees up to 0.50 years. Simulated H. erectus PSCs can be found in all living species by 0.50 years, and the median falls below the human and chimpanzee ranges after 2.5 years. H. erectus ARs are elevated above those of all extant taxa prior to 0.50 years, and after two years they fall out of the human range but are still above ape ranges. A review of evidence for the age at death of Mojokerto supports an estimate of around one year, indicating absolute brain growth rates in the lower half of the human range. These results point to secondary altriciality in H. erectus, implying that key human adaptations for increasing the energy budget of females may have been established by at least 1 Ma.

Skeletal development of hallucal tarsometatarsal joint curvature and angulation in extant apes and modern humans.

The medial cuneiform, namely the curvature and angulation of its distal facet with metatarsal 1, is crucial as a stabilizer in bipedal locomotion and an axis upon which the great toe medially deviates during arboreal locomotion in extant apes. Previous work has shown that facet curvature and angulation in adult dry-bone specimens can distinguish African apes from Homo, and can even distinguish among species of Gorilla. This study provides the first ontogenetic assessment of medial cuneiform curvature and angulation in juvenile (n = 68) and adult specimens (n = 102) using computed tomography in humans and extant ape specimens, including Pongo. Our data find that modern human juveniles initially have a convex and slightly medially oriented osseous surface of the developing medial cuneiform distal facet that flattens and becomes more distally oriented with age. The same pattern (though of a different magnitude) occurs developmentally in the chimpanzee medial cuneiform, but not in Gorilla or Pongo, whose medial cuneiform facet angulation remains unchanged ontogenetically. These data suggest that the medial cuneiform ossifies in a distinguishable pattern between Pongo, Gorilla, Pan, and Homo, which may in part be due to subtle differences in the loading environment at the hallucal tarsometatarsal joint-a finding that has important implications for interpreting fossil medial cuneiforms.

Osteogenic relationship between the lateral plantar process and the peroneal tubercle in the human calcaneus.

The osteogenic relationship between the lateral plantar process and the peroneal tubercle has been an uncertainty for researchers over several decades. While some argue there to be no developmental relationship between these two calcaneal structures, others have suggested that there is an inverse relationship, the lateral plantar process forming from a part of the peroneal tubercle. However, no previous studies have offered quantitative measurements to test these hypotheses. In this study, we measured the size of the peroneal tubercle, retrotrochlear eminence, and the size and area of the lateral plantar process in 73 subjects using magnetic resonance imaging (MRI). Navicular height was measured using weight-bearing radiographs as a measurement of longitudinal arch in 35 of these subjects. Age, body mass, and body mass index (BMI) were also recorded for all subjects. We determined that there was a significant positive correlation between the lateral plantar process and size of the peroneal tubercle, body mass, and BMI. Thus, assertions that there is an inverse relationship between the size of the lateral plantar process and the peroneal tubercle are here unfounded. We also determined there to be a positive correlation between the peroneal tubercle and both the size of the retrotrochlear eminence and the height of the navicular. In conclusion, we relate these novel findings to hominin fossil calcanei and discuss the evolutionary and biomechanical implications.

A shift toward birthing relatively large infants early in human evolution.

It has long been argued that modern human mothers give birth to proportionately larger babies than apes do. Data presented here from human and chimpanzee infant:mother dyads confirm this assertion: humans give birth to infants approximately 6% of their body mass, compared with approximately 3% for chimpanzees, even though the female body weights of the two species are moderately convergent. Carrying a relatively large infant both pre- and postnatally has important ramifications for birthing strategies, social systems, energetics, and locomotion. However, it is not clear when the shift to birthing large infants occurred over the course of human evolution. Here, known and often conserved relationships between adult brain mass, neonatal brain mass, and neonatal body mass in anthropoids are used to estimate birthweights of extinct hominid taxa. These estimates are resampled with direct measurements of fossil postcrania from female hominids, and also compared with estimates of female body mass to assess when human-like infant:mother mass ratios (IMMRs) evolved. The results of this study suggest that 4.4-Myr-old Ardipithecus possessed IMMRs similar to those found in African apes, indicating that a low IMMR is the primitive condition in hominids. Australopithecus females, in contrast, had significantly heavier infants compared with dimensions of the femoral head (n = 7) and ankle (n = 7) than what is found in chimpanzees, and are estimated to have birthed neonates more than 5% of their body mass. Carrying such proportionately large infants may have limited arboreality in Australopithecus females and may have selected for alloparenting behavior earlier in human evolution than previously thought.

Mojokerto revisited: evidence for an intermediate pattern of brain growth in Homo erectus.

Brain development in Homo erectus is a subject of great interest, and the infant calvaria from Mojokerto, Indonesia, has featured prominently in these debates. Some researchers have suggested that the pattern of brain development in H. erectus resembled that of non-human apes, while others argue for a more human-like growth pattern. In this study, we retested hypotheses regarding brain ontogeny in H. erectus using new methods (resampling), and data from additional H. erectus crania. Our results reveal that humans achieve 62% (±10%) and chimpanzees 80% (±9%) of their adult endocranial volume by 0.5-1.5 years of age. Using brain mass data, humans achieve on average 65% and chimpanzees 81% of adult size by 0.5-1.5 years. When compared with adult H. erectus crania (n = 9) from Indonesian sites greater than 1.2 million years old, Mojokerto had reached ∼70% of its adult cranial capacity. Mojokerto thus falls almost directly between the average growth in humans and chimpanzees, and well within the range of both. We therefore suggest that brain development in H. erectus cannot be dichotomized as either ape-like or human-like; it was H. erectus-like. These data indicate that H. erectus may have had a unique developmental pattern that should be considered as an important step along the continuum of brain ontogeny between apes and humans.

Phenotypic plasticity of climbing-related traits in the ankle joint of great apes and rainforest hunter-gatherers.

The "negrito" and African "pygmy" phenotypes are predominately exhibited by hunter-gatherers living in rainforest habitats. Foraging within such habitats is associated with a unique set of locomotor behaviors, most notably habitual vertical climbing during the pursuit of honey, fruit, and game. When performed frequently, this behavior is expected to correlate with developmentally plastic skeletal morphologies that respond to mechanical loading. Using six measurements in the distal tibia and talus that discriminate nonhuman primates by vertical climbing frequency, we tested the prediction that intraspecific variation in this behavior is reflected in the morphology of the ankle joint of habitually climbing human populations. First, to explore the plasticity of climbing-linked morphologies, we made comparisons between chimpanzees, gorillas, and orangutans from wild and captive settings. The analysis revealed significant differences in two climbing-linked traits (anterior expansion of the articular surface of the distal tibia and increased degree of talar wedging), indicating that these traits are sensitive to climbing behavior. However, our analyses did not reveal any signatures of climbing behavior in the ankles of habitually climbing hunter-gatherers. These results suggest that the detection of fine-grained differences in human locomotor behaviors at the ankle joint, particularly those associated with arboreality, may be obscured by the functional demands of terrestrial bipedalism. Accordingly, it may be difficult to use population-level characteristics of ankle morphology to make inferences about the climbing behavior of hominins in the fossil record, even when facultative arborealism is associated with key fitness benefits.

Brief communication: a midtarsal (midfoot) break in the human foot.

The absence of a midtarsal break has long been regarded as a derived feature of the human foot. Humans possess a rigid midfoot that acts as an efficient lever during the propulsive phase of bipedal gait. Non-human primates, in contrast, have a more mobile midfoot that is adaptive for tree climbing. Here, we report plantar pressure and video evidence that a small percentage of modern humans (n = 32/398) possess both elevated lateral midfoot pressures and even exhibit midfoot dorsiflexion characteristic of a midtarsal break. Those humans with a midtarsal break had on average a significantly flatter foot than those without. Midtarsal breakers also had significantly more medial weight transfer (pronation) during the stance phase of gait than those without this midfoot mobility. These data are in accordance with Elftman (Clin Orthop 16 (1960) 41-45) who suggested that pronation aligns the axes of the transverse tarsal joint, permitting elevated midfoot mobility.

Downclimbing and the evolution of ape forelimb morphologies.

The forelimbs of hominoid primates (apes) are decidedly more flexible than those of monkeys, especially at the shoulder, elbow and wrist joints. It is tempting to link the greater mobility of these joints to the functional demands of vertical climbing and below-branch suspension, but field-based kinematic studies have found few differences between chimpanzees and monkeys when comparing forelimb excursion angles during vertical ascent (upclimbing). There is, however, a strong theoretical argument for focusing instead on vertical descent (downclimbing), which motivated us to quantify the effects of climbing directionality on the forelimb kinematics of wild chimpanzees (Pan troglodytes) and sooty mangabeys (Cercocebus atys). We found that the shoulders and elbows of chimpanzees and sooty mangabeys subtended larger joint angles during bouts of downclimbing, and that the magnitude of this difference was greatest among chimpanzees. Our results cast new light on the functional importance of downclimbing, while also burnishing functional hypotheses that emphasize the role of vertical climbing during the evolution of apes, including the human lineage.