RESEARCH DEPARTMENT OF CELL AND DEVELOPMENTAL BIOLOGY

Spoor Palaeolab

Professor Fred Spoor

Contact Details:

Department of Cell and Developmental Biology
University College London (Anatomy Building)
London WC1E 6BT.
Tel. +44 (0)20 7679-4316 (Int. 34316) - Office
Tel. +44 (0)20 7679-4308 (Int. 34308) - Lab
E-mail: f.spoor@ucl.ac.uk

Research

NB: PDFs of articles that tend to be difficult to obtain are available for download below. Please contact me by e-mail if you require PDFs of other publications listed.

Human Evolution

Cranial morphology of Plio-Pleistocene hominins

As a mammalian palaeontologist I study the skull of members of the human (hominin) family tree. This to reconstruct how we as humans evolved over the last six million years. From a broader biological perspective the human head and skull is of particular interest because, unique among mammals, it has been affected by three major phenomena: unprecedented enlargement of the brain, a change to upright posture and gait, and a reduction of the teeth and jaws. Examining how each of these influences is expressed in skull evolution leads me to study the relation between form and function, the developmental (ontogenetic) processes underlying morphological change, and how ancestral form constrains the way a species can adapt to changing external influences.

Most of my work focuses on hominin cranio-dental fossils from the Plio-Pleistocene of eastern Africa, in the context of my affiliation with the Koobi Fora Research Project, directed by Professor Meave Leakey and Dr Louise Leakey (www.kfrp.com). Current projects include the detailed study of the Pliocene fossils from Lomekwi (west of Lake Turkana, Kenya), including the cranium KNM-WT40000 (holotype of Kenyanthropus platyops), and of Plio-Pleistocene fossils found during renewed fieldwork at Koobi Fora and Ileret (east of Lake Turkana, Kenya). A particular focus is on new fossil evidence documenting the radiation of early Homo, a project in collaboration with Dr Susan Antón (New York University), and Professor Christopher Dean (UCL). Supported by the National Geographic Society (USA).

Additional research concerns the Pliocene juvenile Australopithecus afarensis skeleton DIK-1-1 from Dikika, Ethiopia (project directed by Dr Zeresenay Alemseged, Max Planck Institute, Leipzig), and the Pleistocene partial Homo cranium KNM-OG 45500 from Olorgesailie, Kenya (project directed by Dr Rick Potts, Smithsonian Institution).

NB: If you own, or are considering to buy a BoneClones “cast” of the Kenyanthropus cranium KNM-WT 40000, please read this: Spoor F, Leakey LN & Leakey MG (2002) Bone Clones’ “re-creation” of Kenyanthropus. Physical Anthropology 3: 2. [PDF]

Left to right:

Kenyanthropus. platyops cranium KNM-WT 40000 from Lomekwi (Kenya), 3.5 Myr old

Juvenile Australopithecus afarensis skeleton DIK-1-1 from Dikika (Ethiopia), 3.3 Myr old

Homo erectus crania KNM-ER 42700 (small) and OH 9 (large), 1.4-1.6 Myr old

Selected references

Spoor F, Leakey MG, Gathogo PN, Brown FH, Antón SC, McDougall I, Kiarie C, Manthi FK & Leakey LN (2007) Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya. Nature 448, 688-691

Antón SC, Spoor F, Fellmann CD and Swisher III CC (2007) Defining Homo erectus: Size Considered. . In: Handbook of Paleoanthropology Volume III (Henke, Rothe, and Tattersall, eds) Chapter 11, 1655-1693.

Alemseged Z, Spoor F, Kimbel WH, Bobe R, Geraads D, Reed D & Wynn JG. (2006). A juvenile early hominin skeleton from Dikika, Ethiopia. Nature.443: 296-301.

Kupczik K, Spoor F, Pommert A and Dean C (2005). Premolar root number variation in hominoids: genetic polymorphism vs. functional significance. In: “Current Trends in Dental Morphology Research" Zadzinska E (ed), University of Lodz Press, Lodz. pp 257-268. [PDF]

Spoor F, Leakey MG & Leakey LN (2005) Correlation of cranial and mandibular prognathism in extant and fossil hominids. Trans Roy Soc S. Afr. 60: 85-89. [PDF]

Jeffery N, Spoor F. (2004). Ossification and midline shape changes of the human fetal cranial base. Am J Phys Anthrop. 123: 78-90.

Jeffery N & Spoor F (2002) Brain size and the human cranial base: a prenatal perspective. Am. J. Phys. Anthrop. 118: 324-340.

Leakey MG, Spoor F, Brown FH, Gathogo PN, Kiarie C, Leakey LN & McDougall I (2001) New hominin genus from eastern Africa shows diverse middle Pliocene lineages. Nature 410:433.

Spoor F, O'Higgins P, Dean C, Lieberman DE (1999) Anterior sphenoid in modern humans. Nature 397:572.

Spoor, F (1997) Basicranial architecture and relative brain size of Sts 5 (Australopithecus africanus) and other Plio-Pleistocene hominids. South Afr. J. Science 93: 182-187. [PDF]

The Mammalian Inner Ear

Comparative and functional morphology

The bony labyrinth houses the organ of hearing in the cochlea and the sense organs for the perception of movement and spatial orientation in the vestibule and semicircular canals. Of particular interest in my research is the semicircular canal system, which senses head rotations and contributes to the stabilization of gaze and control of locomotion

Comparative studies in collaboration with Professor Alan Walker (Penn State University) show empirically that the arc sizes of the three semicircular canals are significantly larger in mammals that are agile and acrobatic than in species that are more cautious in their locomotion. A main factor underpinning this relationship appears to be that arc size affects the mechanical sensitivity of the canal system. The functional relationship provides the opportunity to assess the locomotor behaviour of extinct taxa, based on their semicircular canals as commonly preserved in fossil crania. Currently fossil primate taxa from the Eocene to the Miocene are being investigated. Supported by the National Science Foundation (USA).

Lateral view of the left bony labyrinths of Galago (left) and Loris (right), reconstructed from high-resolution CT scans. Scale bar is 1 mm. The semicircular canals of the leaping galagid are substantially larger than those of the slow, quadrupedal climbing lorisid (both have similar body size)

Cetaceans (whales and dolphins) have strikingly smaller semicircular canals than other mammals, a phenomenon related to the transition from a terrestrial to a marine environment. In collaboration with Dr Hans Thewissen (North Eastern Ohio College of Medicine, USA) the evolutionary history of this dramatic change is investigated by examining Eocene archaeocete fossils. Supported by the National Science Foundation (USA).

Lateral Side view of the bony labyrinths of, left to right, an agile land mammal (Galago), a land-living early whale from Pakistan (50-million-year-old Ichthyolestes), a marine early whale from India (45-million-year-old Indocetus), and a modern dolphin (Tursiops). The latter two aquatic species have strongly reduced semicircular canals Images reconstructed from CT scans, adjusting for body size

Apart from the link with locomotor behaviour, the bony labyrinth is also affected by shape changes of the surrounding cranial base, and therefore indirectly by changes in brain morphology. Hence, studying the bony labyrinth in fossil hominins is of particular interest because of its association with two key processes of human evolution: the emergence of bipedal locomotion and major brain expansion. Currently my focus is on the bony labyrinth of early Homo and the australopiths.

In collaborative work with Dr Nathan Jeffery (University of Liverpool) we study the fetal development of the primate inner ear, as well as the structural relationship between the subarcuate fossa and the semicircular canals. The latter to assess if spatial constrains of the fossa and its contents (the petrosal lobule of the paraflocculus) are a factor influencing the arc size of the surrounding semicircular canals.

Selected references

Spoor, F., Garland, Th., Krovitz, G., Ryan, T.M., Silcox, M.T. and Walker, A. (2007) The Primate Semicircular Canal System and Locomotion. Proc. Nat. Acad. Sci. 104:10808-10812.

Jeffery N, Spoor F. (2006) The primate subarcuate fossa and its relationship to the semicircular canals part I: prenatal growth. J. Hum. Evol. 51:537-549.

Spoor F, Bajpai S, Hussain ST, Kumar K., Thewissen JGM (2002) Vestibular evidence for the evolution of aquatic behaviour in early cetaceans. Nature 417: 163-166.

Jeffery N, Spoor F. (2004) Prenatal growth and development of the modern human labyrinth. J. Anatomy. 204: 71-92.

Spoor F (2003) The semicircular canal system and locomotor behaviour, with special reference to hominin evolution. Cour. Forsch. Senckenberg. 243: 93-104. [PDF]

Spoor F, Hublin JJ, Braun M & Zonneveld F (2003) The bony labyrinth of Neanderthals. J. Hum Evol. 44: 141-165.

Spoor F and Zonneveld F (1998) Comparative review of the human bony labyrinth. Yearbook of Physical Anthropology 41:211-251.

Spoor F, Stringer C, and Zonneveld F. (1998) Rare temporal bone pathology of the Singa calvaria from Sudan. Am. J. Phys. Anthrop. 107:41-50.

Hublin J-J, Spoor F, Braun M, Zonneveld F and Condemi S. (1996) A late Neanderthal from Arcy-sur-Cure associated with Upper Palaeolithic artefacts. Nature, 381:224-226.

Spoor F, and Leakey M. (1996) Absence of the subarcuate fossa in cercopithecids. J. Human Evol. 31:569-575.

Spoor F, Wood B and Zonneveld F (1994) Implications of early hominid labyrinthine morphology for the evolution of human bipedal locomotion. Nature 369:645-648.

Computed tomography & palaeontology.

Ever since the discovery of x-rays, palaeontology has greatly benefited from radiological techniques such as radiography and, more recently, computed tomography (CT). In the past I have done some research, and have written some reviews on the practical aspects of using CT to study fossils.

Imaging the Homo erectus cranium KNM-WT 15000: lateral radiograph (left), parasagittal CT scan at the level of the right dental row and inner ear (middle), and 3D surface visualization extracted from a stack of CT scans (right). Unlike radiographs, CT scans have the ability to distinguish between fossil bone and the sedimentary matrix in the maxillary sinus (asterisk), and to resolve details such as the root canals of the molars (arrowhead), and structures of the bony labyrinth (arrow).

Selected references

Spoor F, Jeffery N and Zonneveld F. (2000a) Using diagnostic radiology in human evolutionary studies. J. Anatomy 197:61-76.

Spoor F, Jeffery N and Zonneveld F. (2000b) Imaging skeletal growth and evolution. In: Development, Growth and Evolution: implications for the study of the hominid skeleton. (O'Higgins P and Cohn M eds.). Academic Press, London, pp.123-161. [PDF]

Spoor CF, Zonneveld FW and Macho GA (1993) Linear Measurements of Cortical Bone and Dental Enamel by Computed Tomography: Applications and Problems. Am. J. Phys. Anthrop. 91:469-484. [PDF]

Selected blasts from the past.

Hyena locomotion and morphology

For my MSc Biology in the Netherlands I did some work on hyenas: skeletal proportions, gait analysis and muscle dissection of Hyaena hyaena and Crocuta crocuta. To my mild surprise there have been occasional requests over the years for reprints of the resulting articles. These are listed below, with links to PDFs.

Spoor CF and Badoux DM (1989). Descriptive and functional morphology of the locomotory apparatus of the spotted hyena (Crocuta crocuta Erxleben, 1777). Anat. Anz. 168, 261-266. [PDF]

Spoor CF and Badoux DM (1988). Descriptive and functional myology of the back and hindlimb of the striped hyena (Hyaena hyaena L. 1758). Anat. Anz. 167, 313-321. [PDF]

Spoor CF and Badoux DM (1986). Descriptive and functional myology of the neck and forelimb of the striped hyena. Anat. Anz. 161: 375-387. [PDF]

Spoor CF and Belterman Th (1986). Locomotion in Hyaenidae. Contrib. to Zool. 56: 24-28. [PDF]

Spoor CF (1985): Body proportions in Hyaenidae. Anat. Anz. 160: 215-220. [PDF]

Corbeddu Cave (Sardinia)

Between 1982 and 1990 I participated in the palaeontological excavations at Corbeddu Cave, Sardinia, under the direction of the late Paul Sondaar (Institute of Earth Sciences, Utrecht University). Having finished my MSc Biology in 1985 I was not sure what to do for my PhD. With time to kill I opted to describe the human maxilla and temporal bone we had found in pre-Neolithic layers of Corbeddu Cave (~9000 yBP). These specimens, and other evidence, demonstrate that humans co-existed for substantial time with the endemic island fauna of Pleistocene Sardinia. Such co-existence, as opposed to near-instant eradication of endemic island species, is now a phenomenon of particular interest, following the discovery of hominin (“hobbit”) remains as part of the Pleistocene fauna of the island Flores (Indonesia). In any event, this hobby project first triggered my interest in human evolution, solving my dilemma of what direction to take for my PhD.


Left: the 1983 Utrecht University team at Corbeddu Cave. Right: occlusal view of the left maxilla CB85.3013 from Corbeddu Cave, combined with a mirror-imaged cast of the specimen and an extant human molar that fits the wide M¹ alveolus.

Spoor F (1999) The human fossils from Corbeddu Cave, Sardinia: a reappraisal. Deinsea 7:297-302. [PDF]

Spoor CF and Sondaar PY (1986). Human fossils from the endemic island fauna of Sardinia. J. Human Evol. 15: 399 408.

………… Additional esoteric stuff from the past may appear here at a future date.