Steven Churchill

• Duke University

The lower limb of Homo naledi presents a suite of primitive, derived and unique morphological features that pose interesting questions about the nature of bipedal movement in this species. The exceptional representation of all skeletal elements in H. naledi makes it an excellent candidate for biomechanical analysis of gait dynamics using modern kin...

How we teach human genetics matters for social equity. The biology curriculum appears to be a crucial locus of intervention for either reinforcing or undermining students’ racial essentialist views. The Mendelian genetic models dominating textbooks, particularly in combination with racially inflected language sometimes used when teaching about mono...

Studies of human fossils, and the DNA extracted from them, reveal a complex history of interbreeding between various human lineages over the last one hundred thousand years. Of particular interest is the nature of the population interactions between the Neandertals of Ice Age Europe and western Asia and the modern humans that eventually replaced th...

Gi was excavated almost half-a-century ago. Despite no formally published analyses of its lithic and faunal assemblages, the site has been important to discussions about technological development during the Middle Stone Age (MSA). Dating to ~77 ka, ≠Gi has been argued to represent one of the earliest contexts in southern Africa where the spearthrow...

Adaptations of the lower back to bipedalism are frequently discussed but infrequently demonstrated in early fossil hominins. Newly discovered lumbar vertebrae contribute to a near-complete lower back of Malapa Hominin 2 (MH2), offering additional insights into posture and locomotion in Australopithecus sediba . We show that MH2 possessed a lower ba...

Adaptations of the lower back to bipedalism are frequently discussed but infrequently demonstrated in early fossil hominins. Newly discovered lumbar vertebrae contribute to a near-complete lower back of Malapa Hominin 2 (MH2), offering additional insights into posture and locomotion in Australopithecus sediba. We show that MH2 demonstrates a lower...

Homo naledi, a recently discovered hominin species from the Rising Star cave complex in Gauteng Province, South Africa, is a surprising species in more ways than one. The conditions of accumulation, as well as the location of these remains in the cave are intriguing, as is their age of approximately 300,000 years. Likewise, the number of remains as...

A primate's body mass covaries with numerous ecological, physiological, and behavioral characteristics. This versatility and potential to provide insight into an animal's life has made body mass prediction a frequent and important objective in paleoanthropology. In hominin paleontology, the most commonly employed body mass prediction equations (BMP...

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 sam...

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...

Prior to the recovery of Homo naledi from the Dinaledi Chamber of the Rising Star Cave system, the Middle Pleistocene fossil record in Africa was particularly sparse. With the large sample size now available from Dinaledi, the opportunity exists to reassess taxonomically ambiguous teeth unearthed at the nearby site of Sterkfontein. Teeth recovered...

Homo naledi skeletal material described from the Dinaledi Chamber, Rising Star System, in the Cradle of Humankind , South Africa, includes upper limb material with remarkably ape-like morphology occurring in the context of a distinctly modern human-like lower limb, foot, and hand. Here we describe upper limb specimens from a new fossil hominin site...

Humans are thought to exhibit an unusual suite of life history traits relative to other primates, with a longer lifespan, later age at first reproduction, and shorter interbirth interval. These assumptions are key components of popular hypotheses about human life history evolution, but they have yet to be investigated phylogenetically. We applied t...

The lower back is adapted to mobility and stability across mammals and reflects posture and locomotion in the framework of a species’ evolutionary history. Upright bipedalism is one such positional behavior, and due to limited fossil evidence, disagreements exist as to when, how, and in what evolutionary context bipedalism evolved. Here, we describ...

The bones of the pectoral girdle and upper limb are well represented among the fossils thus far recovered from Malapa (Gauteng Province, South Africa). Malapa Hominin 2, a partial skeleton representing an adult female of Australopithecus sediba, preserves a largely complete right pectoral girdle and upper limb, including the most complete scapula y...

While the timing, duration, and nature of the shift is debated, human evolution ultimately involved an increase in body size relative to our Miocene ancestors, and a transition from a more arboreal to a more terrestrial way of life. This is reflected not only in the articular and muscular attachment morphology of the upper and lower limbs, but also...

Australopithecus sediba is known from two partial skeletons, Malapa Hominins 1 and 2 (MH1 and MH2), a juvenile male and an adult female, respectively. Forty-eight elements of the axial skeleton, including vertebrae, ribs, a sternum, and a sacrum, are known from MH1 and MH2. Here, we describe these ~2.0 Ma fossils and provide raw data and plots of s...

The exceptionally well-preserved Neanderthal skeleton discovered in October 1993 within the Lamalunga cave near Altamura (Puglia, Italy) has been recently dated to a late Middle Pleistocene chronology, bracketed between 128.2 and 187.0 ka. Although the skeleton is still sealed in situ, in 2009 and 2015 a large part of its fragmentary right scapula...

We report here preliminary results from four seasons of excavation at the rockshelter of Riparo Bombrini (2002–2005). Three markedly separate horizons were uncovered: the deepest, comprising Levels M1-7, yielded abundant Mousterian lithics and faunal remains. A second macro-unit, corresponding to Levels MS1-2, is only a few decimeters thick and is...

The upper limb of Homo naledi has previously been described from fragmentary material discovered in the Dinaledi Chamber in the Rising Star cave system, Cradle of Humankind, South Africa. Recent work at Rising Star has led to the recovery of additional fossil material attributed to H. naledi from a new fossil site within the system called the Lesed...

Objectives: Predicting body mass is a frequent objective of several anthropological subdisciplines, but there are few published methods for predicting body mass in immature humans. Because most reference samples are composed of adults, predicting body mass outside the range of adults requires extrapolation, which may reduce the accuracy of predict...

Abstract The enigmatic Homo naledi has many special characteristics. This geologically recent species is characterized by a mosaic of derived and archaic characters. Today, 12 clavicles and 15 scapulas have been discovered, coming from the cave chambers Dinaledi (site 101) and Lesedi (also known as Site 102). The scapulas are very fragmentary and a...

In the hominin fossil record, pelvic remains are sparse and are difficult to attribute taxonomically when they are not directly associated with craniodental material. Here we describe the pelvic remains from the Dinaledi Chamber in the Rising Star cave system, Cradle of Humankind, South Africa, which has produced hominin fossils of a new species, H...

New fossil discoveries and new analyses increasingly blur the lines between Australopithecus and Homo, changing scientific ideas about the transition between the two genera. The concept of the genus itself remains an unsettled issue, though recent fossil discoveries and theoretical advances, alongside developments in phylogenetic reconstruction and...

Postcranial measurements. DOI: http://dx.doi.org/10.7554/eLife.24232.047

Canonical variates analysis of carpal morphology. DOI: http://dx.doi.org/10.7554/eLife.24232.048

Traits of the LES1 cranium in comparison to H. naledi and other hominin species. DOI: http://dx.doi.org/10.7554/eLife.24232.045

Cranial and mandibular measurements. DOI: http://dx.doi.org/10.7554/eLife.24232.046

Taphonomic observations by specimen from the Lesedi Chamber. DOI: http://dx.doi.org/10.7554/eLife.24232.049

ELife digest Species of ancient humans and the extinct relatives of our ancestors are typically described from a limited number of fossils. However, this was not the case with Homo naledi. More than 1500 fossils representing at least 15 individuals of this species were unearthed from the Rising Star cave system in South Africa between 2013 and 2014...

Body mass is an ecologically and biomechanically important variable in the study of hominin biology. Regression equations derived from recent human samples allow for the reasonable prediction of body mass of later, more human-like, and generally larger hominins from hip joint dimensions, but potential differences in hip biomechanics across hominin...

The evolution of the hominin pelvis is generally seen as involving two broad stages: the establishment of bipedal pelvic morphology by the mid-Pliocene (or earlier), followed by architectural changes necessary to enlarge the birth canal in response to increased encephalization in Pleistocene members of the genus Homo. Pelvic and proximal femoral mo...

H. naledi shows a mosaic morphological pattern with several derived (Homo-like) features of the skull, hands and feet, and primitive (australopith- like) features in the ribcage, shoulder, and pelvis. This pattern reflects a morphology that might be expected of a hominin at the evolutionary transition between Australopithecus and Homo. Two thoracic...

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...

The evolutionary transition from an ape-like to human-like upper extremity occurred in the context of a behavioral shift from an upper limb predominantly involved in locomotion to one adapted for manipulation. Selection for overarm throwing and endurance running is thought to have further shaped modern human shoulder girdle morphology and its posit...

We describe the earliest evidence for neoplastic disease in the hominin lineage. This is reported from the type specimen of the extinct hominin Australopithecus sediba from Malapa, South Africa, dated to 1.98 million years ago. The affected individual was male and developmentally equivalent to a human child of 12 to 13 years of age. A penetrating l...

Lithic projectile points are a universal component of the hunting tool kits of archeologically- and historically-known foragers. Recent experimental work with ballistic gelatin target s has shown that lithic-tipped projectiles do not have a marked penetration advantage over those with simple sharpened wooden points, leading to the suggestion that i...

Adult human foragers expend roughly 30–60 kcal per km in unburdened walking at optimal speeds.1,2 In the context of foraging rounds and residential moves, they may routinely travel distances of 50–70 km per week, often while carrying loads.3 Movement on the landscape, then, is arguably the single most expensive item in the activity budgets of hunte...

Rising Star Cave is located in the Cradle of Humankind World Heritage site near Krugersdorp in South Africa. In November 2013 and March 2014 more than 1500 hominin fossil elements attributed to a new species, Homo naledi, were recovered and catalogued representing at least a dozen individuals. Only 20 out of 206 bones in the human body are not in t...

In this preliminary reconstruction of Homo naledi’s gait we begin with the null hypothesis that it walked similarly to modern humans, as the overall anatomy of this extinct hominin’s lower limb, especially its foot, is mostly modern human-like. We note the following characters as modern-like: dorsally-canting metatarsophalangeal joints facilitating...

It is largely held that the transition from Australopithecus to the genus Homo involved a shift in overall body proportions, including a relative lengthening of the lower limb, though the degree to which hominin limb proportions changed and the proximate causes thereof remain contentious topics. The overall poor preservation of most articular surfa...

In the hominin fossil record, pelvic remains are sparse and are difficult to attribute taxonomically when they are not directly associated with crania. Here we introduce the pelvic remains from the Dinaledi Chamber in the Rising Star cave system, Cradle of Humankind, South Africa which belong to the newly discovered species, Homo naledi. Though thi...

The recently announced species, Homo naledi (Rising Star Cave, Cradle of Humankind, South Africa), combines an australopith-like cranial capacity with dental characteristics more akin to early Homo. Its postcranium is similarly mosaic; although the hand, foot, and lower limb share many derived characteristics, the H. naledi shoulder girdle and uppe...

Presenting first results of a comparative analysis of the Australopithecus sediba costal material using 3D geometric morphometrics..

The thorax of Australopithecus sediba is hypothesized to follow a mosaic evolutionary pattern showing a greater similarity with Australopithecus afarensis and great apes in the upper thorax and a greater similarity with humans at the lower thorax and waist (Berger et al. 2011; Schmid et al. 2013). This study presents first results of a comparative...

We describe the axial skeletal material recovered from the Dinaledi chamber of Rising Star cave that, together with some pelvis and shoulder remains, document the complicated nature of trunk evolution. The axial material includes two near-complete lower thoracic vertebrae found in articulation with an 11th rib, the proximal portion of a 12th rib, a...

Modern humans are characterized by a highly specialized foot that reflects our obligate bipedalism. Our understanding of hominin foot evolution is, although, hindered by a paucity of well-associated remains. Here we describe the foot of Homo naledi from Dinaledi Chamber, South Africa, using 107 pedal elements, including one nearly-complete adult fo...

A nearly complete right hand of an adult hominin was recovered from the Rising Star cave system, South Africa. Based on associated hominin material, the bones of this hand are attributed to Homo naledi. This hand reveals a long, robust thumb and derived wrist morphology that is shared with Neandertals and modern humans, and considered adaptive for...

Electron microprobe analyses of spots in fragments of samples UW101-SO-31, UW101-SO-34, UW101-SO-39 and DB-1. Note that in each of the tables totals below 100% reflect volatile content or porosity of sample, or both. DOI: http://dx.doi.org/10.7554/eLife.09561.018

Summary table listing surface modifications on all morphologically informative specimens. A total of 559 bone and dental specimens were examined for surface modifications. This sample includes all of the larger specimens and most of the complete elements in the collection, from both surface and excavation contexts. At low magnification (7×) most of...

Traits of H. naledi and comparative species. DOI: http://dx.doi.org/10.7554/eLife.09560.029

Holotype and paratype specimens and referred materials. DOI: http://dx.doi.org/10.7554/eLife.09560.028

Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. na...

Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. na...

Neandertals and Homo sapiens are known to differ in scapular glenoid fossa morphology. Functional explanations may be appropriate for certain aspects of glenoid fossa morphology; however other factors - e.g., allometry, evolutionary development - must be addressed before functional morphology is considered. Using three dimensional geometric morphom...

Sex determination is critical for developing the biological profile of unidentified skeletal remains. When more commonly used elements (os coxa, cranium) for sexing are not available, methods utilizing other skeletal elements are needed. This study aims to assess the degree of sexual dimorphism of the lumbar vertebrae and develop discriminant funct...

For us, the experience of reading Steve Churchill's book Thin on the Ground: Neandertal Biology, Archeology, and Ecology was like that of reading The Origin of Species for the first time. In both Churchill's and Darwin's books, the reader is led carefully and meticulously through a beautifully organized presentation of all the evidence bearing on a...

Inferences about Neandertal home range sizes have historically been reconstructed with reference to lithic raw material transport distances. Here we use data on northern latitude social carnivores to predict home range sizes for Neandertal groups. Given that Neandertals must have relied heavily on animal protein and fat in the plant food-poor envir...

This chapter provides a sketch of the major habitats and climates in which Neandertals found themselves, as a background against which to explore their capture and use of energy. Recent work on North American Pleistocene faunas indicates that mammalian communities are continually and unpredictably emergent, and that many Late Pleistocene faunal com...

This chapter focuses on the activity costs-the caloric costs of finding and procuring food, water, shelter, raw materials for tools, and fuel ? that were added on to the Neandertal caloric budget. It reviews the evidence for Neandertal activity levels in an effort to better estimate the caloric costs of subsistence and other activities in their ene...

This chapter focuses on active predation by which Neandertals procured their prey and zooarcheological and human fossil evidence, to delineate Neandertal hunting strategies and tactics. This is central to efforts to better understand the energetic costs of foraging, some of the limitations that operated on rates of energy capture, and the overall e...

As an approach to understanding Neandertal reproductive ecology and its effects on demography, this chapter explores the inferences that can be made about Neandertal socioecology, and then once again pick up the thread of Neandertal energetics. The chapter begins by asking what can be said about the size of Neandertal social groups ? both in terms...

Increasingly, archeologists and cognitive psychologists are examining the role of cultural, historical, ecological, and demographic factors in both cumulative technological evolution (CTE) and the persistent expression of symbolic behavior. The chapters in this book advance two mutually compatible arguments as to why the Neandertals lived at such l...

The Neandertals are associated with a rich and well-studied archeological record, comprised of items of material culture, dietary residues, and domestic features. This chapter provides a brief overview of Neandertal material culture generally, its focus is on the adaptively-significant aspects of Neandertal technology. While the time of origin of t...

This chapter explores the implications of both types of competition to Neandertal ecology, with an eye towards defining the niche space occupied by Neandertals vis-à-vis other carnivores, assessing habitat use constraints experienced by Neandertals because of the presence of other large-bodied carnivores, and estimating population densities of the...

Recent isotopic and trace element analyses reveal that the European Neandertals were deriving most all of their dietary protein from animal sources, suggesting that their diets were dominated by meat and other animal tissues. The chapter addresses questions of diet (what items were included, what mix of foods from different trophic levels were cons...

This chapter first examines climatic reconstructions for the glacial cycles between 240–30 Ka BP. It then reviews the evidence for cold adaptation in Neandertals including the inferences that make about morphological adaptation from their skeletal morphology and about behavioral adaptations from the archeological record. This review undertakes with...

Any attempt to define the Neandertals and delineate their evolutionary history soon bumps into several thorny issues, including (1) the appropriate choice of species concept when dealing with fossil taxa, (2) the most appropriate methods of taxonomic diagnosis, and (3) how best to reconstruct phylogenetic relationships between paleospecies. Thus de...

While the situation in Middle Pleistocene Africa may not have been as severe as it was for the energy-strapped, relatively meat-dependent Neandertals, the combination of high energy demands and aggressive interactions with carnivores may have been limiting factors on population growth for both groups. By late Middle Pleistocene times modern humans...

Body size is the single most important determinant of an organism's biology, impacting everything from physiological processes to ecological relationships. Given a diet that was rich in animal tissues, Neandertal body size would have been central to predatory success, as well as to the outcome of competitive interactions with carnivore competitors....

The past 200,000 years of human cultural evolution have witnessed the persistent establishment of behaviors involving innovation, planning depth, and abstract and symbolic thought, or what has been called “behavioral modernity.” Demographic models based on increased human population density from the late Pleistocene onward have been increasingly in...