Fred Spoor

New hominin genus from eastern Africa shows diverse middle Pliocene lineages

Most interpretations of early hominin phylogeny recognize a single early to middle Pliocene ancestral lineage, best represented by Australopithecus afarensis, which gave rise to a radiation of taxa in the late Pliocene. Here we report on new fossils discovered west of Lake Turkana, Kenya, which differ markedly from those of contemporary A. afarensis, indicating that hominin taxonomic diversity extended back, well into the middle Pliocene. A 3.5 Myr-old cranium, showing a unique combination of derived facial and primitive neurocranial features, is assigned to a new genus of hominin. These findings point to an early diet-driven adaptive radiation, provide new insight on the association of hominin craniodental features, and have implications for our understanding of Plio–Pleistocene hominin phylogeny.

A juvenile early hominin skeleton from Dikika, Ethiopia

Understanding changes in ontogenetic development is central to the study of human evolution. With the exception of Neanderthals, the growth patterns of fossil hominins have not been studied comprehensively because the fossil record currently lacks specimens that document both cranial and postcranial development at young ontogenetic stages. Here we describe a well-preserved 3.3-million-year-old juvenile partial skeleton of Australopithecus afarensis discovered in the Dikika research area of Ethiopia. The skull of the approximately three-year-old presumed female shows that most features diagnostic of the species are evident even at this early stage of development. The find includes many previously unknown skeletal elements from the Pliocene hominin record, including a hyoid bone that has a typical African ape morphology. The foot and other evidence from the lower limb provide clear evidence for bipedal locomotion, but the gorilla-like scapula and long and curved manual phalanges raise new questions about the importance of arboreal behaviour in the A. afarensis locomotor repertoire.

The primate semicircular canal system and locomotion.

The semicircular canal system of vertebrates helps coordinate body movements, including stabilization of gaze during locomotion. Quantitative phylogenetically informed analysis of the radius of curvature of the three semicircular canals in 91 extant and recently extinct primate species and 119 other mammalian taxa provide support for the hypothesis that canal size varies in relation to the jerkiness of head motion during locomotion. Primate and other mammalian species studied here that are agile and have fast, jerky locomotion have significantly larger canals relative to body mass than those that move more cautiously.

Vestibular evidence for the evolution of aquatic behaviour in early cetaceans

Early cetaceans evolved from terrestrial quadrupeds to obligate swimmers, a change that is traditionally studied by functional analysis of the postcranial skeleton. Here we assess the evolution of cetacean locomotor behaviour from an independent perspective by looking at the semicircular canal system, one of the main sense organs involved in neural control of locomotion. Extant cetaceans are found to be unique in that their canal arc size, corrected for body mass, is approximately three times smaller than in other mammals. This reduces the sensitivity of the canal system, most plausibly to match the fast body rotations that characterize cetacean behaviour. Eocene fossils show that the new sensory regime, incompatible with terrestrial competence, developed quickly and early in cetacean evolution, as soon as the taxa are associated with marine environments. Dedicated agile swimming of cetaceans thus appeared to have originated as a rapid and fundamental shift in locomotion rather than as the gradual transition suggested by postcranial evidence. We hypothesize that the unparalleled modification of the semicircular canal system represented a key ‘point of no return’ event in early cetacean evolution, leading to full independence from life on land.

Comparative review of the human bony labyrinth

The bony labyrinth inside the petrous part of the temporal bone houses the organs of hearing and balance. Being functionally linked with sensory control of body movements and located in a part of the basicranium that is closely associated with the brain, this structure is of great interest in the study of human evolutionary history. However, few aspects of its morphology have been studied in nonhuman primates. This review compares the bony labyrinth of humans with that of the great apes and 37 other primate species based on data newly acquired with computed tomography combined with previous descriptions. With body mass taken into account, consistent differences are found between the size of the semicircular canals in humans, the great apes, and other primates. In particular, the arcs of the anterior and posterior canals are larger in humans than in the African apes. The functional implications of semicircular canal dimensions for registering angular head motion are evaluated in relation to locomotor behavior. Biophysical models, comparative studies, and some neurophysiological experiments all support a link between semicircular canal size and agility, or more specifically the frequency contents of natural head movements, but the evidence on the exact nature of this link is ambiguous. It is concluded that any link between the characteristic dimensions of the human canals and locomotion will be more complex than a simple association with the broad categories of quadrupedal vs. bipedal behavior. The functionally important planar orientations of the semicircular canals are similar in humans and the African apes as well as in many other species. In contrast, other aspects of the human labyrinth differ markedly in shape, following a pattern that seems to reflect the characteristic architecture of the human basicranium. Indeed, it is found that labyrinthine and basicranial shape are interspecifically correlated in the sample, and in most respects the human morphology is consistent with the general trend among primate species. Differences in brain growth and development are proposed as the predominant factor underlying both the unique shape of the human labyrinth as well as the interspecific labyrintho-basicranial correlations.

The bony labyrinth of Neanderthals

This paper presents a comprehensive comparative analysis of the Neanderthal bony labyrinth, a structure located inside the petrous temporal bone. Fifteen Neanderthal specimens are compared with a Holocene human sample, as well as with a small number of European Middle Pleistocene hominins, and early anatomically modern and European Upper Palaeolithic humans. Compared with Holocene humans the bony labyrinth of Neanderthals can be characterized by an anterior semicircular canal arc which is smaller in absolute and relative size, is relatively narrow, and shows more torsion. The posterior semicircular canal arc is smaller in absolute and relative size as well, it is more circular in shape, and is positioned more inferiorly relative to the lateral canal plane. The lateral semicircular canal arc is absolutely and relatively larger. Finally, the Neanderthal ampullar line is more vertically inclined relative to the planar orientation of the lateral canal. The European Upper Palaeolithic and early modern humans are most similar, although not fully identical to Holocene humans in labyrinthine morphology. The European Middle Pleistocene hominins show the typical semicircular canal morphology of Neanderthals, with the exception of the arc shape and inferiorly position of the posterior canal and the strongly inclined ampullar line. The marked difference between the labyrinths of Neanderthals and modern humans can be used to assess the phylogenetic affinities of fragmentary temporal bone fossils. However, this application is limited by a degree of overlap between the morphologies. The typical shape of the Neanderthal labyrinth appears to mirror aspects of the surrounding petrous pyramid, and both may follow from the phylogenetic impact of Neanderthal brain morphology moulding the shape of the posterior cranial fossa. The functionally important arc sizes of the Neanderthal semicircular canals may reflect a pattern of head movements different from that of modern humans, possibly related to aspects of locomotor behaviour and the kinematic properties of their head and neck.

New fossils from Koobi Fora in northern Kenya confirm taxonomic diversity in early Homo

Since its discovery in 1972 (ref. 1), the cranium KNM-ER 1470 has been at the centre of the debate over the number of species of early Homo present in the early Pleistocene epoch of eastern Africa. KNM-ER 1470 stands out among other specimens attributed to early Homo because of its larger size, and its flat and subnasally orthognathic face with anteriorly placed maxillary zygomatic roots. This singular morphology and the incomplete preservation of the fossil have led to different views as to whether KNM-ER 1470 can be accommodated within a single species of early Homo that is highly variable because of sexual, geographical and temporal factors, or whether it provides evidence of species diversity marked by differences in cranial size and facial or masticatory adaptation. Here we report on three newly discovered fossils, aged between 1.78 and 1.95 million years (Myr) old, that clarify the anatomy and taxonomic status of KNM-ER 1470. KNM-ER 62000, a well-preserved face of a late juvenile hominin, closely resembles KNM-ER 1470 but is notably smaller. It preserves previously unknown morphology, including moderately sized, mesiodistally long postcanine teeth. The nearly complete mandible KNM-ER 60000 and mandibular fragment KNM-ER 62003 have a dental arcade that is short anteroposteriorly and flat across the front, with small incisors; these features are consistent with the arcade morphology of KNM-ER 1470 and KNM-ER 62000. The new fossils confirm the presence of two contemporary species of early Homo, in addition to Homo erectus, in the early Pleistocene of eastern Africa.

Prenatal growth and development of the modern human labyrinth

The modern human bony labyrinth is morphologically distinct from that of all other primates, showing derived features linked with vestibular function and the overall shape of the cranial base. However, little is known of how this unique morphology emerges prenatally. This study examines in detail the developing fetal human labyrinth, both to document this basic aspect of cranial biology, and more specifically, to gain insight into the ontogenetic basis of its phylogenetically derived morphology. Forty‐one post‐mortem human fetuses, ranging from 9 to 29 weeks gestation, were investigated with high‐resolution magnetic resonance imaging. Quantitative analyses of the labyrinthine morphology revealed a number of interesting age‐related trends. In addition, our findings show that: (1) the prenatal labyrinth attains an adult equivalent size between 17 and 19 weeks gestation; (2) within the period investigated, shape changes to all or most of the labyrinth cease after the 17–19‐week size maturation point or after the otic capsule ossifies; (3) fetal cochlea development correlates with the surrounding petrosal morphology, but not with the midline basicranium; (4) gestational age‐related rotations of the ampullae and cochlea relative to the lateral canal, and posterior canal torsion are similar to documented phylogenetic trends whereas other trends remain distinct. Findings are discussed in terms of the ontogenetic processes and mechanisms that most likely led, in part, to the emergence of the phylogenetically derived adult modern human labyrinth.

Climate-related variation of the human nasal cavity

The nasal cavity is essential for humidifying and warming the air before it reaches the sensitive lungs. Because humans inhabit environments that can be seen as extreme from the perspective of respiratory function, nasal cavity shape is expected to show climatic adaptation. This study examines the relationship between modern human variation in the morphology of the nasal cavity and the climatic factors of temperature and vapor pressure, and tests the hypothesis that within increasingly demanding environments (colder and drier), nasal cavities will show features that enhance turbulence and air-wall contact to improve conditioning of the air. We use three-dimensional geometric morphometrics methods and multivariate statistics to model and analyze the shape of the bony nasal cavity of 10 modern human population samples from five climatic groups. We report significant correlations between nasal cavity shape and climatic variables of both temperature and humidity. Variation in nasal cavity shape is correlated with a cline from cold-dry climates to hot-humid climates, with a separate temperature and vapor pressure effect. The bony nasal cavity appears mostly associated with temperature, and the nasopharynx with humidity. The observed climate-related shape changes are functionally consistent with an increase in contact between air and mucosal tissue in cold-dry climates through greater turbulence during inspiration and a higher surface-to-volume ratio in the upper nasal cavity.

Using diagnostic radiology in human evolutionary studies

This paper reviews the application of medical imaging and associated computer graphics techniques to the study of human evolutionary history, with an emphasis on basic concepts and on the advantages and limitations of each method. Following a short discussion of plain film radiography and pluridirectional tomography, the principles of computed tomography (CT) and magnetic resonance imaging (MRI) and their role in the investigation of extant and fossil morphology are considered in more detail. The second half of the paper deals with techniques of 3‐dimensional visualisation based on CT and MRI and with quantitative analysis of digital images.