Digital Conservation and Reconstruction of 'Fossilized Behaviour': the evolution of the human foot as revealed by ancient footprint trails

Lead Research Organisation: Bournemouth University
Department Name: Sch of Applied Sciences

Abstract

Fossil footbones are rarely found associated with identifiable skeletons, and are often fragmentary; and when we do find a partial foot of one of our ancient ancestors it has usually been badly chewed. In addition such fossil rarely give definite indications of the way our early ancestors walked, as they act through a nested series of complicated soft tissues, from ligaments, out to the skin, and therefore the bones interact only remotely with the ground on which we walk. On the other hand, the footprints and trackways which were left when our distant ancestors walked across soft ground are the closest we can come to 'fossilized behaviour', as they are direct records of the forces we apply to the ground to balance ourselves and propel our walking. They are therefore potentially excellent evidence of the evolution of human walking. However, until recently we have lacked the tools with which to unlock their scientific potential. For example, the functional significance of the famous Laetoli footprints made some 3.75 million years ago by 'Lucy' and her relatives has been argued about for over 30 years, one scientist contradicting another, on the basis of features of individual prints - and in one case mistaking the footprint of a hare which walked across the human ancestor's footprints as the print of its big toe. We need methods which will tell us what are the common features of fossil trackways; identifying their 'central tendency' so that we can eliminate bias. This isn't easy, since footprints, being made by soft tissue, have no easily recognisable landmark points. One of the two groups involved in this project work on the mechanics of walking, using computer-simulation techniques, and engaging in some 'lateral thinking', found that methods used to analyze the distribution of chemical patterns in the brain are ideal for comparing footprints. The other group are specialists in finding and excavating fossil footprints, and recently discovered an exciting new set of prints in Kenya, about half the age of the Laetoli prints and made by the first members of our own 'genus': Homo. They used advanced laser-scanning techniques to record the prints in three-dimensional detail, as they had previously done for those at Laetoli. It's very likely that in the next three years they will find even more new footprint trails. Human walking works like a metronome, saving energy as the body swings forward over the foot which contacts the ground, which can then be used to power the next stride. But because of our tall, thin build, our walking is unstable from side to side, and our hip muscles need to work to counter our tendency to fall over sideways when one leg takes over support from the other. Short, squatter animals like penguins are more stable from side to side, and can actually save pendulum-energy sideways on, by their 'waddling' gait. Distant human ancestors like Lucy had a similar squat and stable build - did they save energy the same way? It is likely that the footprints at Laetoli may contain the answer. Comparing them with the new footprints of early Homo should tell us a lot about how Lucy's flexible foot changed into a stiffer one which could push-off hard enough to let us walk or run long distances nearly effortlessly. To do so, we need not only to make sophisticated computer models of walking and footprint formation, which can recreate balance and energy-saving mechanisms in these early human ancestors, and relate them to foot forces and footprint form, but to do physical experiments in soft mud and ash which will tie these models into the real world. But we must also look for more footprints, which can fill in some of the details of the changes we are studying. Thus, working together, we have the skills, tools and evidence to interpret the evolution of walking at a crucial time period, the transition between an early biped that probably spent some time in the trees, and a striding long-distance walker.

Publications

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Bates KT (2013) Does footprint depth correlate with foot motion and pressure? in Journal of the Royal Society, Interface

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Bates KT (2013) The evolution of compliance in the human lateral mid-foot. in Proceedings. Biological sciences

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Bennett M (2014) Tracks made by swimming Hippopotami: An example from Koobi Fora (Turkana Basin, Kenya) in Palaeogeography, Palaeoclimatology, Palaeoecology

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Bennett M (2010) Probable Neolithic footprints preserved in inter-tidal peat at Kenfig, South Wales (UK) in Proceedings of the Geologists' Association

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Bennett M (2016) Footprints and human evolution: Homeostasis in foot function? in Palaeogeography, Palaeoclimatology, Palaeoecology

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Gierlinski G (2017) Possible hominin footprints from the late Miocene (c. 5.7 Ma) of Crete? in Proceedings of the Geologists' Association

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Larsen HJ (2021) Recovery of 3D footwear impressions using a range of different techniques. in Journal of forensic sciences

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Matthew Bennett (Author) (2010) Footprints of the Kuiseb Delta, Namibia in The Digging Stick

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Matthew Bennett (Author) (2010) Tracking our Ancestors in NERC Planet Earth

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Matthew R Bennett (2014) Digitising Human Tracks: challenges and Developments in Geological Society of America: Annual Meeting and Exposition

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Morse SA (2013) Holocene footprints in Namibia: the influence of substrate on footprint variability. in American journal of physical anthropology

 
Description The research set out to use fossil footprints - fossilised locomotion - to examine the potential difference in foot anatomy and function across the crucial and important evolution transition from the Australopiths to Homo. A particular focus was the tracks (1.6 Ma) from Ileret (Kenya) believed to be made by Homo erectus and documented by Bennett in 2009 in Science. These tracks were combined with those of Laetoli (Australopithecus afarensis; 3.66 Ma) and modern unshod tracks from Namibia documented for the first time as part of this research. Tracks were also studied from Argentina and the Sefton Coast in the UK.
The research involved an innovative combination of modelling (both substrate and biomechanics), fieldwork, laboratory analysis using pressure treadmills and modern analogue experiments. Innovative analytical methods in the form of pedobariographic Statistical Parametric Mapping (pSPM) was used to create mean tracks from whole populations, co-register these means and then compare them statistically. It became clear early in the research that understanding the influence of substrate and inter-site differences was going to be critical to success. Consequently much focus, especially by Bennett, was placed on understanding the influence of sedimentary substrate in mediating footprint morphology. This focus led to some of the most innovative and exciting results of the research. A Holocene footprint site in Namibia, documented in detail for the first time, was used to explore the influence of substrate, developing conceptual field models for the role of substrate in modifying and preserving track morphology (Morse et al. 2013; Bennett 2014). In addition, using modelling and modern analogue data the group explored the relationship between depth and plantar pressure. The potential correlation between these two variable is fundamental to the interpretation of vertebrate tracks and while the relationship held in general it was found to be complex in detail with significant implications for the interpretation of tracks of palaeo-anthropological significance (Bates et al. 2013a,b).
The team made an initial comparison between the Laetoli tracks from in Tanzania are 3.65 million year using scans obtained by Bennett from the National Museum of Kenya and both shod and unshod modern humans. The innovative analytical approach based on pSPM allowed statistical comparison to be made for the first time (Crompton et al. 2014). The main conclusion here was that the anatomy and foot function of the Laetoli track maker was essentially modern and not dramatically different from tracks made by Homo sapiens. This implies a significant period of homo stasis in foot function across this major evolutionary transition. Difficulties in isolating good tracks from Ileret, due to substrate influence, limited the team's ability to broaden this analysis. Despite this means have been created for the Ileret tracks and comparison with those of modern humans and the Laetoli track maker confirm this period of homo stasis. Further work is on-going to improve the registration and statistical comparison of these footprint populations, including the development of new analytical software. Further publications are due shortly. Irrespective of this, the research developed new analytical tools, including software, for the study of human tracks establishing a clear methodology/benchmark by which new sites should be examined like those from Happisburgh (UK) documented at the start of 2014. Much of the research completed under this NERC award is summarised in the book by Bennett and Morse (2014) which sets out in form of a definitive manual what can and cannot be inferred from human footprints and how their future study should be approached.



Footprints & Substrate: Namibia



Working at a Holocene human footprint site in the Namib Sand Sea, south of Walvis Bay, Namibia the team has explored intra-trail footprint variability associated with small variations in substrate properties using 'whole foot' analytical techniques. High levels of intra-trail variability have been found as a result of variations in grain-size, depositional moisture content and the degree of sediment disturbance, all of which determine the bearing capacity of the substrate. The method, model and results presented here are essential in the interpretation of sites of paleoanthropological significance, such as recently reported from Ileret (1.5 Ma, Kenya; Bennett et al., Science, 2009). Footprint typology varies with bearing capacity such that firm substrates show limited impressions associated with areas of peak plantar pressure, while softer substrates are associated with deep prints with narrow heels and reduced medial longitudinal arches. Substrates of medium bearing capacity give displacement rims and proximal movement of sediment, which obscures the true form of the medial longitudinal arch. A simple conceptual model has been developed which summarizes these conclusions.



Sources: Morse et al. (2013) American Journal of Physical Anthropology, in press.







Geoconservation of Footprint Sites & Methods of Digital Field Recording



Human footprints provide some of the most publically emotive and tangible evidence of our ancestors. To the scientific community they provide evidence of stature, presence, behaviour and in the case of early hominins potential evidence with respect to the evolution of gait. Many footprint sites are at risk from rapid erosion, including the Ileret footprints in northern Kenya which are second only in age to those at Laetoli (Tanzania). Unlithified, soft-sediment footprint sites such these pose a significant geoconservation challenge. The team has explored the conservation and preservation options available at such sites and concluded that to 'record and digitally rescue' them provides the only viable approach. Key to such strategies is the increasing availability of three-dimensional data capture either via optical laser scanning and/or digital photogrammetry. Within the discipline there is a developing schism between those that favour one approach over the other and a requirement from geoconservationists and the scientific community for some form of objective appraisal of these alternatives is necessary. Consequently the team has evaluated these alternative approaches and the role they can play in a 'record and digitally rescue' conservation strategy. Using modern footprint data, digital models have been created via optical laser scanning and compared to those generated by state-of-the-art photogrammetry. Both methods give comparable although subtly different results. This data is evaluated alongside a review of field deployment issues to provide guidance to the community with respect to the factors which need to be considered in digital conservation of human/hominin footprints. This methodological comparison is of value to end users both in geoconservation and forensic science.



Source: Bennett et al. (2013) Plos One, in press.







Footprint Depth & Biomechanics



Footprints provide a direct source of evidence about locomotor biomechanics in extinct vertebrates. One of the principal suppositions underpinning biomechanical inferences is that footprint geometry correlates with dynamic foot pressure, which, in turn, is linked with overall limb motion of the trackmaker. The team has performed the first quantitative test of this longstanding assumption, using topological statistical analysis of plantar pressures and experimental and computer-simulated footprints. In computer-simulated footprints, the relative distribution of depth differed from the distribution of both peak and pressure impulse in all simulations. Analysis of footprint samples with common loading inputs and similar depths reveals that only shallow footprints lack significant topological differences between depth and pressure distributions. Topological comparison of plantar pressures and experimental beach footprints demonstrates that geometry is highly dependent on overall print depth; deeper footprints are characterized by greater relative forefoot, and particularly toe, depth than shallow footprints. The highlighted difference between 'shallow' and 'deep' footprints clearly emphasizes the need to understand variation in foot mechanics across different degrees of substrate compliance. Overall, our results indicate that extreme caution is required when applying the 'depth equals pressure' paradigm to hominin footprints, and by extension, those of other extant and extinct tetrapods.



Source: Bates et al. (2013) Journal of the Royal Society Interface, doi: 10.1098/rsif.2013.0009.
Exploitation Route We recognize three main end users of our research: (1) forensic crime investigators; (2) security processes and personal identification; and (3) medical and care industry in relation to biomechanics in elderly populations - trip and fall prevention - and the early diagnosis of foot conditions such as diabetic foot and osteoarthritis. The software (pSPM) which is protected an international patent is a key tool in footprint and foot pressure analysis. Other software for three-dimensional visualization - Foot Processor - has also been developed. We are adopted so far a range of routes including: (1) academic conferences and workshops; (2) public outreach; and (3) briefings and collaborative workshops with two police forces.
Sectors Digital/Communication/Information Technologies (including Software),Education,Environment,Pharmaceuticals and Medical Biotechnology,Security and Diplomacy,Other

 
Description Our work has been used widely by the scientific community and forms a benchmark for the analysis of human tracks. We are currently exploring forensic applications and developing software applications to make this possible. This has lead to a NERC Innovation Award in 2015.
First Year Of Impact 2014
Sector Digital/Communication/Information Technologies (including Software),Security and Diplomacy
Impact Types Policy & public services

 
Description Forensic Footwear 
Organisation Home Office
Department Home Office Scientific Development Branch
Country United Kingdom 
Sector Public 
PI Contribution Both organisations are project partners of the NERC Innovation Award and are actively helping to develop the forensic software being developed.
Collaborator Contribution Contacts, know-how and support.
Impact The software development is on going, due for completion in June 2016.
Start Year 2015
 
Description Forensic Footwear 
Organisation National Crime Agency
Country United Kingdom 
Sector Public 
PI Contribution Both organisations are project partners of the NERC Innovation Award and are actively helping to develop the forensic software being developed.
Collaborator Contribution Contacts, know-how and support.
Impact The software development is on going, due for completion in June 2016.
Start Year 2015
 
Title Foot Processor 
Description It is a tool for the analysis of three-dimensional track data, essentially the visualisation of XYZ point clouds. It is freeware based on a free Mathworks compiler. It is has widespread use for researchers working in vertebrate track studies. 
Type Of Technology Software 
Year Produced 2013 
Open Source License? Yes  
Impact The software is now widely used and downloaded by the research community. We are currently re-writing (in Python) and enhancing this software and hope to develop forensic applications for it. 
URL http://footprints.bournemouth.ac.uk/
 
Title Foot Viewer 
Description The software allows multiple ASC/CSV XYZ files to be viewed simultaneously. It is a research tool aimed at the vertebrate track community 
Type Of Technology Software 
Year Produced 2013 
Open Source License? Yes  
Impact The software is widely used by the community and regularly downloaded. 
URL http://footprints.bournemouth.ac.uk/
 
Title Track Transformer 
Description This is a tool for the comparison of vertebrate tracks. 
Type Of Technology Software 
Year Produced 2015 
Open Source License? Yes  
Impact This software is part of the code being developed in the current NERC Innovation award and was central to the publication Bennett et al. (2016) on the Laetoli tracks in Tanzania. 
 
Description 14th Congress of the PanAfrican Archaeological Association of Prehistory and Related Studies 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Participants in your research and patient groups
Results and Impact The project team delivered a number of presentations at this conference including: PRESERVING THE IMPOSSIBLE: CONSERVATION OF SOFT-SEDIMENT HOMININ FOOTPRINT SITES; THE HOMININ FOOTPRINT SITES OF TURKANA: COMPARISON OF ANIMAL TRACKS AND PALAEOENVIRONMENTS; and HOLOCENE FOOTPRINTS IN NAMIBIA: ARCHAEOLOGY AND FOOTPRINT LABORATORY. All presentations were well received and have lead to a number of approaches around future collaboration.

Several vertebrate track researchers have been in touch with data and offers of future collaboration.
Year(s) Of Engagement Activity 2014
 
Description Ancient footprint tramples on theory of human evolution The Times 04/09/2017  
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact New paper interview and publications (both print and online) following publication of paper in Proceeding of the Geologists' Association in 2017
Year(s) Of Engagement Activity 2017
 
Description Ancient footprints reveal lifestyle of children 700,000 years age; The Independent (Web) 15/02/2018  
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact News paper interview and report following publication of paper in Scientific Reports on: Ancient footprints reveal lifestyle of children 700,000 years age
Year(s) Of Engagement Activity 2017
URL https://www.independent.co.uk/news/science/ancient-footprints-children-lifestyle-700000-years-ago-a8...
 
Description Discovery Magazine January 2018 - Top 100 Discoveries/Stories in 2017; Cretan Footprints were #8 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Discovery reported the Cretan Footprints after publication in September 2017 and listed in a third article in 2018 as #8 in importance in their top 100 stories.
Year(s) Of Engagement Activity 2018
 
Description Foot Lab 2014 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach Regional
Primary Audience Public/other audiences
Results and Impact The Foot Lab ran as part of the BU Festival of Learning in 2014. The aim was to attract participants to leave their tracks for science. It was both a data gathering and an outreach activity aimed at public engagement.

There was a large amount of interested and over 150 volunteers participated in the activity.
Year(s) Of Engagement Activity 2014
 
Description Museum of Natural History New Mexico 2014 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact The aim was to engage with them around our research and demonstrate its applicability in museum settings. The intervention lead to multiple offers of collaboration and also to the acquisition of track data from the museum archive.

Established an on-going route for future collaboration.
Year(s) Of Engagement Activity 2014
 
Description Newly discovered 6m-year-old Cretan footprints stolen - finder writes about how we must protect precious sites, The Conversation 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Article in The Conversation following theft of the Cretan Footprints. This has been read over 11,500 times.
Year(s) Of Engagement Activity 2017
URL https://theconversation.com/newly-discovered-6m-year-old-cretan-footprints-stolen-finder-writes-abou...
 
Description Our controversial footprint discovery suggests human-like creatures may have roamed Crete nearly 6m years ago, The Conversation 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Article entitled 'Our controversial footprint discovery suggests human-like creatures may have roamed Crete nearly 6m years ago' in The Conversation following publication in September 2017 of the paper on Cretan Footprints. It has been read over 105,000 times.
Year(s) Of Engagement Activity 2017
URL https://theconversation.com/our-controversial-footprint-discovery-suggests-human-like-creatures-may-...
 
Description Thief legs it with ancient footprints The Times 16/09/2017  
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Interview and news report (Online, Print both UK and Irish Times) on the theft of the Cretan Footprints.
Year(s) Of Engagement Activity 2017
 
Description Understanding footprints: forensic applications 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Geographic Reach National
Primary Audience Professional Practitioners
Results and Impact Briefing for Metropolitan Police which led to discussions about how to develop our work for practical uses. These discussions have continued and are the potential subject of an Innovation Award by the applicants.

Strong interest from participants leading to continued contact and discussion
Year(s) Of Engagement Activity 2012
 
Description What ancient footprints can tell us about what it was like to be a child in prehistoric times, The Conversation February 2018 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Article in the conversation entitled 'What ancient footprints can tell us about what it was like to be a child in prehistoric times ' following publication of our Scientific Reports paper in February 2018 so far this has been read 31,000 times.
Year(s) Of Engagement Activity 2018
URL https://theconversation.com/what-ancient-footprints-can-tell-us-about-what-it-was-like-to-be-a-child...