School of Anthropology & Conservation

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Dr Tracy Kivell

Reader in Biological Anthropology

Primate locomotion; skeletal morphology; origin and evolution of human bipedalism and hand use.


profile image for Dr Tracy Kivell

School roles and responsibilities

Director of the Animal Postcranial Evolution (APE) Lab.

Academic background

I am a palaeoanthropologist who studies the functional morphology of the wrist and hand in extant and fossil primates. My research focuses on extant and fossil apes, including fossil hominins, to further our understanding of the origin of human bipedalism and hand use throughout our evolutionary history. I aim to understand the relationship between bone shape and function of the hand through analyses of ontogeny, internal (trabecular and cortical) bone structure, and the biomechanics of primate locomotion. My current research interests include:

  1. Functional morphology of the Australopithecus sediba hand
  2. Trabecular and cortical bone structure in extant and fossil primate hands to better understand locomotion and tool-use in early hominins
  3. Biomechanical analyses of terrestrial and arboreal locomotion in apes
  4. Functional morphology of fossil hand bones of Miocene hominoids


I received my PhD from the University of Toronto, working on the developmental morphology of the ape wrist and the origin of human bipedalism.  Before coming to the University of Kent, I was a Research Associate, teaching human gross anatomy at Duke University (2007-2009) and was postdoctoral Junior Researcher at the Max Planck Institute for Evolutionary Anthropology (2009-2013). I have participated in palaeontological field excavations at Miocene sites in Hungary as well as Plio-Pleistocene sites in South Africa.

Tracy is also the Director of the Animal Postcranial Evolution (APE) Laboratory

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Also view these in the Kent Academic Repository

Book section
Kivell, T. et al. (2016). Introduction. in: Kivell, T. L. et al. eds. The Evolution of the Primate Hand: Anatomical, Developmental, Functional, and Paleontological Evidence. Springer, pp. 1-3. Available at:
Kivell, T. (2016). The Primate Wrist. in: The Evolution of the Primate Hand: Anatomical, Developmental, Functional, and Paleontological Evidence. Springer.
Edited book
Kivell, T.L. et al. eds. (2016). The Evolution of the Primate Hand: Anatomical, Developmental, Functional, and Paleontological Evidence. [Online]. Springer. Available at:
Stephens, N. et al. (2016). Trabecular architecture in the thumb of Pan and Homo: implications for investigating hand use, loading, and hand preference in the fossil record. American Journal of Physical Anthropology [Online]:1-17. Available at:
Kivell, T. (2016). A review of trabecular bone functional adaptation: what have we learned from trabecular analyses in extant hominoids and what can we apply to fossils? Journal of Anatomy [Online] 228:569-594. Available at:
Schoonaert, K. et al. (2016). Gait characteristics and spatio-temporal variables of climbing in bonobos (Pan paniscus). American Journal of Primatology [Online]:1-13. Available at:
Behringer, V. et al. (2016). Within Arm’s Reach: Measuring Forearm Length to Assess Growth Patterns in Captive Bonobos and Chimpanzees. American Journal of Physical Anthropology [Online]:1-7. Available at:
Kivell, T. (2015). Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa. eLife [Online]:1-37. Available at:
Showing 8 of 38 total publications in KAR. [See all in KAR]



A full list of publications is included in my CV.

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Bone structure of a Chimp hand

I occasionally teach on undergraduate (e.g. SE 302) and graduate (e.g. SE856)  modules, but will not be teaching full time until September 2020.

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Science cover featuring 2011 research paper

GRASP  Evolution of the human hand: Grasping trees and tools (funded by European Research Council Starting Grant 2014-2019)  See “Research Projects”or click here

Director of research on Australopithecus sediba hand fossils: Investigation of Au. sediba and Rising Star hand remains from South Africa in collaboration with Lee Berger and Job Kibii (University of the Witwatersrand) and Steve Churchill (Duke University).

Fossil hominin and hominoid hand use: Comparative investigation of fossil hominin (australopiths to Neandertals) and Miocene hominoid hand remains using morphometric and micro-CT data. This research aims to shed light on locomotor and tool-use behaviours throughout the evolution of the human lineage. Collaborators: Matthew Skinner (UCL), Steve Churchill (Duke University), David Begun (University of Toronto).

Functional signals in trabecular and cortical bone structure: A comparative investigation of internal bony morphology of the primate hand to assess variation in joint loading patterns and how this reflects differences in locomotor and manipulative behaviours. Collaborators: Matthew Skinner (UCL), Dieter Pahr and Thomas Gross (Vienna University of Technology), Huynh Nguyen and Jean-Jacques Hublin (Max Planck Institute-EVA) and Richard Lazenby (University of Northern British Columbia).

Biomechanics of primate locomotion: Investigation of variation in biomechanics of terrestrial and arboreal locomotion in apes as well as other primates, including the aye aye, in both captive and natural settings. Collaborators: Daniel Schmitt and Brian Hare (Duke University) and Roshna Wunderlich (James Madison University).

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Orangutan trabecular bone

I can offer supervision of PhD and MA/MSc research within any of my areas of interest – functional morphology of the postcranial skeleton, including external and internal (using microCT data) bony morphology – with a particular focus on the upper limb.

Current PhD students

Principal Supervisor

Johanna NeufussHand use and posture during locomotor and non-locomotor behaviours in wild, habituated gorillas, chimpanzees and bonobos


Chris Dunmore"Skeletal form and function of the primate hand"
Leoni Georgiou "Functional morphology of the hip and knee joints in apes and humans"
Zewdi Tsegai
(Max Planck Institute for Evolutionary Anthropology)
Nicholas Stephens (Max Planck Institute for Evolutionary Anthropology)

Past Masters students

Ann-Marie SchillingTrabecular bone structure in the primate wrist

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Aye Aye as featured on the cover of the Journal of Experimental Biology


Physical Anthropology Women’s Mentoring Network (PA WMN), an organization that brings together more senior women in biological anthropology with younger women seeking advice as they transition from their PhD to a postdoc, to their first academic positions and through the tenure process, balancing work, research and family. We organize mentoring events at the American Association of Physical Anthropology (AAPA) conference each year, but offer mentoring year round. I currently sit on the Steering Committee and was Chair of PA WMN in 2011-2012.

Symposium Organizer (with Prof. Evie Vereecke) of the “Primates and their hands: behavioural, experimental and morphological perspectives on the primate hand”.  European Federation for Primatology, Antwerp, Belgium, Sept. 2013.

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I am available to provide commentary or discussion on human or nonhuman primate evolution, primate locomotion, or skeletal morphology and evolution.




  • 2013  BBC4 documentary “Hands and feet”, aired February 2014
  • 2012  Three-part documentary, “Your Inner Fish” Windfall Films, DVD due to be
    released 2014
  • 2009  The Daily Planet on Discovery Channel, Canada: “The World is Just Awesome Week”, Episode 13, September 14th: “Unraveling the secrets behind these not-so-cute creatures”


Popular magazines and websites             

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Last Updated: 05/07/2016