Hominin Footprints Research Articles

A Late Pleistocene hominin footprint site on the North African coast of Morocco

Footprints represent a relevant vestige providing direct information on the biology, locomotion, and behaviour of the individuals who left them. However, the spatiotemporal distribution of hominin footprints is heterogeneous, particularly in North Africa, where no footprint sites were known before the Holocene. This region is important in the evolution of hominins. It notably includes the earliest currently known Homo sapiens (Jebel Irhoud) and the oldest and richest African Middle Stone Age hominin sites. In this fragmented ichnological record, we report the discovery of 85 human footprints on a Late Pleistocene now indurated beach surface of about 2800 m2 at Larache (Northwest coast of Morocco). The wide range of sizes of the footprints suggests that several individuals from different age groups made the tracks while moving landward and seaward across a semi-dissipative bar-trough sandy beach foreshore. A geological investigation and an optically stimulated luminescence dating of a rock sample extracted from the tracksite places this hominin footprint surface at 90.3 ± 7.6 ka (MIS 5, Late Pleistocene). The Larache footprints are, therefore, the oldest attributed to Homo sapiens in Northern Africa and the Southern Mediterranean.

Neanderthal footprints in the “Matalascañas trampled surface” (SW Spain): new OSL dating and Mousterian lithic industry

In the Huelva Coast of SW Spain erosion by recent marine storms revealed the presence of a paleosol where an extensive tracksite known as “Matalascañas Trampled Surface” (MTS) has been documented. The MTS includes tracks and trackways of large species of mammals, along with bird trace fossils, invertebrate burrows and root traces. Within this record, the presence of several hominin footprints and trackways stands out. Despite previous uncertainties about the producer of these footprints, new OSL age of 151 ± 11 ka secures their attribution to Neanderthals, the only hominins known to have been present in the Iberian Peninsula during the MIS6-5 transition. Moreover, typical Mousterian lithic industry with Levallois knapping was found associated with the ichnological record. This lithic industry is characterized by the selection of raw materials from outcrops in a short-distance range to the tracksite. The general characteristics of the lithics are derived both from the nature of the raw material and from the nature of the site itself, which cannot be seen as a settlement, but rather as a place of passage for fauna, including Neanderthals, where a few human individuals performed short-term activities, such as food procurement and/or meat processing.

Fossil footprints at the late Lower Paleolithic site of Schöningen (Germany): A new line of research to reconstruct animal and hominin paleoecology

The ca. 300 ka Paleolithic sites of Schöningen in northern Germany yielded a number of localities with archeological and paleontological remains representing a rich paleoenvironmental record of the late Middle Pleistocene in northern Europe. An important line of research focused on the ichnology of two localities: Schöningen 13 I-Fs2 and Schöningen 13 II-2 Untere Berme. Here we present the first detailed study of these fossil footprints, which provides insights on Schöningen's paleoenvironment and a snapshot of the mammals once living in the area. Herds of elephants and other species of herbivores congregated along the muddy shores of a paleolake during birch, pine and grass-rich woodland phases. In addition, three potential hominin footprints, probably belonging to a late Homo heidelbergensis, are recorded at Schöningen 13 II-2 Untere Berme. This confirms the complementary potential of ichnology in reconstructing a reliable picture of prehistoric sites.

Laetoli: The Oldest Known Hominin Footprints in Volcanic Ash

Hominin footprints are rare in eastern Africa and known from the Laetoli (Tanzania), Ileret (Kenya), and Melka Kunture (Ethiopia) areas. The prints were made by Australopithecus afarensis, Homo erectus, and Homo heidelbergensis. Their study is an important source of information regarding hominin body size, anatomy, positional behavior, and locomotion biomechanics. The most-known and best-studied examples are the 3.66-Ma Australopithecus afarensis footprint trackways at Laetoli, which represent the oldest known record of hominin bipedalism in Africa. The footprints occur in a volcanic tuff sequence, which was originally deposited as melilite nephelinite ash. Recent excavations show that this valuable paleoanthropological site is slowly disappearing as a result of surface diagenetic processes. Preservation of the footprints is essential and urgently needed.

New dating of the Matalascañas footprints provides new evidence of the Middle Pleistocene (MIS 9-8) hominin paleoecology in southern Europe

Hominin footprints were recently discovered at Matalascañas (Huelva; South of Iberian Peninsula). They were dated thanks to a previous study in deposits of the Asperillo cliff to 106 ± 19 ka, Upper Pleistocene, making Neandertals the most likely track-makers. In this paper, we report new Optically Stimulated Luminescence dating that places the hominin footprints surface in the range of 295.8 ± 17 ka (MIS 9-MIS 8 transition, Middle Pleistocene). This new age implies that the possible track-makers are individuals more likely from the Neandertal evolutionary lineage. Regardless of the taxon attributed to the Matalascañas footprints, they supplement the existing partial fossil record for the European Middle Pleistocene Hominins being notably the first palaeoanthropological evidence (hominin skeleton or footprints) from the MIS 9 and MIS 8 transition discovered in the Iberian Peninsula, a moment of climatic evolution from warm to cool. Thus, the Matalascañas footprints represent a crucial record for understanding human occupations in Europe in the Pleistocene.

Fossil footprints and what they mean for hominin paleobiology.

Hominin footprints have not traditionally played prominent roles in paleoanthropological studies, aside from the famous 3.66 Ma footprints discovered at Laetoli, Tanzania in the late 1970s. This contrasts with the importance of trace fossils (ichnology) in the broader field of paleontology. Lack of attention to hominin footprints can probably be explained by perceptions that these are exceptionally rare and "curiosities" rather than sources of data that yield insights on par with skeletal fossils or artifacts. In recent years, however, discoveries of hominin footprints have surged in frequency, shining important new light on anatomy, locomotion, behaviors, and environments from a wide variety of times and places. Here, we discuss why these data are often overlooked and consider whether they are as "rare" as previously assumed. We review new ways footprint data are being used to address questions about hominin paleobiology, and we outline key opportunities for future research in hominin ichnology.

From footprint morphometrics to the stature of fossil hominins: A common but uncertain estimate

Hominin footprints are a particular remain in paleoanthropology representing brief moments of life of extinct individuals. Footprints not only provide information on the locomotor behavior of fossil taxa but also on their body characteristics such as their stature. This stature is usually estimated from the length of the footprints based on the well-known foot length to stature ratio. However, footprint length does not result only from the foot length but of a combination of factors. Therefore, it is necessary to investigate the relationships between footprint length and stature of individuals using experimental approaches. Secondly, recent discoveries of fossil footprints have led to the estimation of statures from isolated footprints. However, such estimates may be biased because of the intraindividual morphometric variation of the footprints. Moreover, footprints may also be incomplete making it impossible to measure the length and therefore the estimation of a stature. The search for relationships between stature and other morphometric variables is therefore necessary to have the most accurate picture possible of the individuals who left these tracks. In this context, this article reports the results of an experimental study that aims to determine the relationships between the stature of individuals and different morphometric variables and to quantify the intraindividual variation of each variable. Thus, 21 morphometric variables were measured on a total of 175 experimental footprints left by 20 individuals in an experimental area composed of loose sand. Statistical analyses show that footprint lengths are not only the variables most correlated with stature but also those with the least intraindividual variation. However, estimation of stature from footprints left by fossil hominins is subject to three types of uncertainties: residuals from linear regression, intraindividual variation that can be particularly large in soft substrates, and the application of relationships defined on modern populations to fossil taxa.

Une vue d’ensemble sur Melka Kunture, grand complexe de sites pléistocènes dans la vallée supérieure de l’Awash (Éthiopie)

Melka Kunture is a cluster of Pleistocene sites, extending over ⁓100 km2 between 2000 and 2200m asl, in the upper Awash Valley of Ethiopia. Starting around 2 million-years ago, the archaeological sequence includes sites with lithic productions of the Oldowan, Early Acheulean, middle Acheulean, final Acheulean, Early Middle Stone Age, Middle Stone Age and Late Stone Age. All over the Pleistocene, the climate was rainy and cooler than at the lower elevations of the Rift Valley, allowing the development of Afromontane vegetation. Hippopotamuses are ubiquitous and dominant in terms of biomass, but Alcelaphini are well represented, notably with genus Connochaetes and genus Damaliscus. Hominin fossils have been discovered in association with the Oldowan, the Early Acheulean, the middle Acheulean and the Early Middle Stone Age. Animal tracks and hominin footprints have also been documented, the latter ones in layers dated between 1.2 and 0.7 million-years.

Age constraints for the Trachilos footprints from Crete

We present an updated time frame for the 30 m thick late Miocene sedimentary Trachilos section from the island of Crete that contains the potentially oldest hominin footprints. The section is characterized by normal magnetic polarity. New and published foraminifera biostratigraphy results suggest an age of the section within the Mediterranean biozone MMi13d, younger than ~ 6.4 Ma. Calcareous nannoplankton data from sediments exposed near Trachilos and belonging to the same sub-basin indicate deposition during calcareous nannofossil biozone CN9bB, between 6.023 and 6.727 Ma. By integrating the magneto- and biostratigraphic data we correlate the Trachilos section with normal polarity Chron C3An.1n, between 6.272 and 6.023 Ma. Using cyclostratigraphic data based on magnetic susceptibility, we constrain the Trachilos footprints age at ~ 6.05 Ma, roughly 0.35 Ma older than previously thought. Some uncertainty remains related to an inaccessible interval of ~ 8 m section and the possibility that the normal polarity might represent the slightly older Chron C3An.2n. Sediment accumulation rate and biostratigraphic arguments, however, stand against these points and favor a deposition during Chron C3An.1n.

The Le Rozel footprints: snapshots of Neandertal groups in the Late Pleistocene. A combined morphometric and experimental approach

Hominin footprints represent brief moments of life and provide access to the locomotor and biological features of the individuals who left them. Because of the particular time scale that they represent and the information they provide, they make it possible to approach the size and composition of hominin groups in a different way from the methods used with the more common archaeological artefacts and osteological assemblages. Since 2012, several hundred Neandertal footprints have been discovered at Le Rozel (Normandy, France), within 5 stratigraphic sub-units dated to about 80,000 years ago. Analyses of the ichnological assemblage discovered at Le Rozel between 2012 and 2017 have identified 257 footprints, 132 of which have been digitized in 3D. This represents the largest ichnological assemblage attributed to Neandertals to date. Morphometric analyses of the 3D models of the footprints, combined with an experimental study under substrate conditions similar to those at Le Rozel, have quantified the size and composition of the groups that left them. The footprints from the most ichnologically dense stratigraphic sub-unit were left by a small group and represent 90% of children and adolescents. These results not only enhance our understanding of the Palaeolithic occupations at Le Rozel but also provide new methods and data for assessing the size and composition of Neandertal groups.

Body mass estimation from footprint size in hominins

Although many studies relating stature to foot length have been carried out, the relationship between foot size and body mass remains poorly understood. Here we investigate this relationship in 193 adult and 50 juvenile habitually unshod/minimally shod individuals from five different populations—Machiguenga, Daasanach, Pumé, Hadzabe, and Samoans—varying greatly in body size and shape. Body mass is highly correlated with foot size, and can be predicted from foot area (maximum length × breadth) in the combined sample with an average error of about 10%. However, comparisons among populations indicate that body shape, as represented by the body mass index (BMI), has a significant effect on foot size proportions, with higher BMI samples exhibiting relatively smaller feet. Thus, we also derive equations for estimating body mass from both foot size and BMI, with BMI in footprint samples taken as an average value for a taxon or population, estimated independently from skeletal remains. Techniques are also developed for estimating body mass in juveniles, who have relatively larger feet than adults, and for converting between foot and footprint size. Sample applications are given for five Pliocene through Holocene hominin footprint samples from Laetoli (Australopithecus afarensis), Ileret (probable Homo erectus), Happisburgh (possible Homo antecessor), Le Rozel (archaic Homo sapiens), and Barcin Höyük (H. sapiens). Body mass estimates for Homo footprint samples appear reasonable when compared to skeletal estimates for related samples. However, estimates for the Laetoli footprint sample using the new formulae appear to be too high when compared to skeletal estimates for A. afarensis. Based on the proportions of A.L. 288-1, this is apparently a result of relatively large feet in this taxon. A different method using a ratio between body mass and foot area in A.L. 288-1 provides estimates more concordant with skeletal estimates and should be used for A. afarensis.

The morphological affinity of the Early Pleistocene footprints from Happisburgh, England, with other footprints of Pliocene, Pleistocene, and Holocene age

Fossil hominin footprints provide a direct source of evidence of locomotor behavior and allow inference of other biological data such as anthropometrics. Many recent comparative analyses of hominin footprints have used 3D analytical methods to assess their morphological affinities, comparing tracks from different locations and/or time periods. However, environmental conditions can sometimes preclude 3D digital capture, as was the case at Happisburgh (England) in 2013. Consequently, we use here a 2D geometric morphometric approach to investigate the evolutionary context of the Happisburgh tracks. The comparative sample of hominin tracks comes from eight localities that span a broad temporal range from the Pliocene to Late Holocene. The results show disparity in the shapes of tracks ascribed to hominins from the Pliocene (presumably Australopithecus afarensis), Pleistocene (presumably Homo erectus and Homo antecessor), and Holocene (Homo sapiens). Three distinct morphological differences are apparent between time samples: changes in adduction of the hallux, changes in the shape and position of the medial longitudinal arch impression, and apparent changes in foot proportions. Linear dimensions classified the potential H. antecessor tracks from Happisburgh as being most similar to the presumed H. erectus prints from Ileret. We demonstrate using 2D geometric morphometric methods and linear dimensions that the Happisburgh tracks are morphologically similar to other presumed Homo tracks and differ from the Laetoli footprints. The probable functional implications of these results fit well with previous comparative analyses of hominin tracks at other sites.

Snapshots of human anatomy, locomotion, and behavior from Late Pleistocene footprints at Engare Sero, Tanzania

Fossil hominin footprints preserve data on a remarkably short time scale compared to most other fossil evidence, offering snapshots of organisms in their immediate ecological and behavioral contexts. Here, we report on our excavations and analyses of more than 400 Late Pleistocene human footprints from Engare Sero, Tanzania. The site represents the largest assemblage of footprints currently known from the human fossil record in Africa. Speed estimates show that the trackways reflect both walking and running behaviors. Estimates of group composition suggest that these footprints were made by a mixed-sex and mixed-age group, but one that consisted of mostly adult females. One group of similarly-oriented trackways was attributed to 14 adult females who walked together at the same pace, with only two adult males and one juvenile accompanying them. In the context of modern ethnographic data, we suggest that these trackways may capture a unique snapshot of cooperative and sexually divided foraging behavior in Late Pleistocene humans.

The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France)

Footprints represent a unique snapshot of hominin life. They provide information on the size and composition of groups that differs from osteological and archeological remains, whose contemporaneity is difficult to establish. We report here on the discovery of 257 footprints dated to 80,000 y from the Paleolithic site at Le Rozel (Normandy, France), which represent the largest known Neandertal ichnological assemblage to date. We investigate the size and composition of a track-maker group from this large set by developing a morphometric method based on experimental footprints. Our analyses indicate that the footprints were made by a small group comprising different age classes, from early childhood to adult, with a majority of children. The Le Rozel footprints thus provide direct evidence for the size and composition of a Neandertal social group.

Local people’s interpretations of the hominin footprints at Laetoli, Tanzania

ABSTRACTIn 1976 and 2014, scientists reported hominin footprints in Pliocene volcanic deposits at Laetoli, north-eastern Tanzania. They hypothesized that the footprints belong to the extinct hominin species Australopithecus afarensis. Not only have the footprints shaped scientists’ understanding of human biological evolution, but they have also placed Laetoli and Tanzania on the world’s paleoanthropological studies map. However, there have been different interpretations of who made these footprints. Opinions of the scientists on the footprints are widely documented, yet narratives of Maasai (local people living around Laetoli) have never been investigated. This paper documents the perceptions of Maasai on what these footprints are, and to whom they belong. We report that concepts related to the nature and genesis of the footprints and who made them form part of the Maasai’s history. Finally, we suggest that for sustainable footprints preservation, local narratives about footprints be integrated into scientific interpretations of the site.

Using differential structure-from-motion photogrammetry to quantify erosion at the Engare Sero footprint site, Tanzania

Hominin footprint sites are excellent sources of data that provide insights into early human physiology, anatomy, and social structures. They are also potential tourist attractions that are often situated in relatively under-developed parts of the world. Unfortunately, many footprint sites are also located in high energy environments and/or are pressed into poorly indurated sediments, which make them highly susceptible to erosion. This paper proposes a non-invasive and low-cost method employing Structure-from-Motion photogrammetry to quantify erosion rates in the absence of permanent ground control points. Using point cloud comparison algorithms between data collected at different times, it is possible to quantitatively analyze the locations, volumes, and rates of material loss. We applied this technique to several footprints within the Engare Sero footprint site in northern Tanzania to assess erosional change between 2010 and 2017. Our comparisons show that prints are vertically eroding at average rates ranging from 0.10 to 0.17 mm/yr with some localized areas experiencing much higher rates over shorter durations. We identify three primary modes of erosion: 1) flaking, 2) abrasion, and 3) boring of holes. Erosion appears to have occurred episodically with major events, such as flooding, separated by periods of relative stability. The methods presented here are valuable for paleoanthropologists to better understand how footprint erosion might adversely affect inferences regarding print-makers, and they are valuable for decision-makers, who can create conservation plans to better protect and maximize the utility of known hominin footprint sites.

Comments on possible Miocene hominin footprints

Gierlinski et al. (2017) report on what they interpret to be Miocene hominin footprints near the seaside village of Trachilos in western Crete. We review the case made by the authors that these ichnites represent bona-fide footprints, and their conclusion that they were made by bipedal hominins. Gierlinski et al.'s study demonstrates a number of problems with data presentation, e.g. a) substrates corresponding to measured prints are not clearly specified, b) no explanation is given for how prints were identified when the authors' own criteria for print identification were not met, c) no consistent morphological detail among prints is provided that could identify them as originating from the same or a similar agent, or one with bilateral symmetry, d) alternative agents that could have produced the prints are not explored, e) no explanation is given as to how their multivariate analyses of print outlines deals with missing data and why it uses non-homologous landmarks, etc. The evidence they present, therefore, is insufficient to support their arguments and conclusions. We remain unconvinced the ichnites are bona-fide footprints, let alone hominin footprints, but discuss some of the criteria employed for distinguishing and recognizing an early hominin footprint.

Using experimental biomechanics to reconstruct the evolution of hominin locomotor postures

Debates over the evolution of hominin bipedalism, a defining human characteristic, often revolve around whether early bipeds walked more like humans, with energetically economical extended hind limbs, or more like apes with flexed hind limbs. Here, we detail experimental data from humans and chimpanzees that links anatomy with lower limb posture and the energy costs of walking. Using these anatomical links, we examine the fossil record to determine when adaptations evolved that allowed for extended limb bipedalism.To complement anatomical studies of hominin fossils, we apply evolutionary biomechanics to the 3.6 million year old hominin footprints at Laetoli, Tanzania, which represent the earliest direct evidence of hominin bipedalism. We compare footprints from the original Lateoli Site G (n=8) and the newly described Site S (n=8) with experimentally generated footprints from modern humans (n=8 subjects) made using both extended and flexed limb, or Bent‐Knee, Bent‐Hip (BKBH) biomechanics. In our modern human sample, footprints generated during BKBH walking leave deeper toe relative to heel impressions compared with footprints generated during normal, extended limb walking (linear mixed effects model: p<0.001). The difference in proportional toe depth (calculated as log10[maximum fore‐foot depth/maximum heel depth]) is linked to the mechanical consequences of BKBH walking. During walking, the center of pressure (COP; the point of ground force application) moves from the heel at touchdown to the forefoot during toe‐off, and the forces applied to the ground determine, to some degree, the depth of the impression under the COP. As the COP travels past the mid‐foot the human heel rises due to the presence of a stiff longitudinal arch, and forces following heel rise deform the substrate under the toes. From experimentally generated force‐plate records, we demonstrate that the COP passes the mid‐foot earlier in the step in BKBH walking, leading to larger forces under the toes and relatively deeper impressions.Using linear mixed effects models, we show that prints from Site G and the newly described prints from Site S at Laetoli are most consistent with weight transfer patterns from extended limb biomechanics. Proportional toe depths do not differ significantly between the Laetoli sites (G vs. S: p=0.998), and do not differ between Laetoli and modern humans walking with an extended limb gait (p=0.195). However, the print morphology at Laetoli does differ significantly from those made by humans walking with a flexed limb (p<0.001) due to the uniquely deep toe depressions that occur in humans walking with this more ape‐like gait.Thus, using experimental data from living humans, we interpret morphology from both the Laetoli footprints and from early hominin skeletons to propose extended limb bipedal posture evolved by 3.6 Ma. These results represent the earliest direct evidence of kinematically human‐like bipedalism currently known. Thus, our results, suggest that selection acted to increase the energy economy of bipedalism early in human evolution and that efficient extended limb bipedalism evolved long before the appearance of the genus Homo.Support or Funding InformationFunded by the L.S.B. Leakey Foundation and the University of ArizonaThis abstract is from the Experimental Biology 2018 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.

Introduction to special issue: Body mass estimation — Methodological issues and fossil applications

Although many studies relating stature to foot length have been carried out, the relationship between foot size and body mass remains poorly understood. Here we investigate this relationship in 193 adult and 50 juvenile habitually unshod/minimally shod individuals from five different populations—Machiguenga, Daasanach, Pumé, Hadzabe, and Samoans—varying greatly in body size and shape. Body mass is highly correlated with foot size, and can be predicted from foot area (maximum length × breadth) in the combined sample with an average error of about 10%. However, comparisons among populations indicate that body shape, as represented by the body mass index (BMI), has a significant effect on foot size proportions, with higher BMI samples exhibiting relatively smaller feet. Thus, we also derive equations for estimating body mass from both foot size and BMI, with BMI in footprint samples taken as an average value for a taxon or population, estimated independently from skeletal remains. Techniques are also developed for estimating body mass in juveniles, who have relatively larger feet than adults, and for converting between foot and footprint size. Sample applications are given for five Pliocene through Holocene hominin footprint samples from Laetoli (Australopithecus afarensis), Ileret (probable Homo erectus), Happisburgh (possible Homo antecessor), Le Rozel (archaic Homo sapiens), and Barcin Höyük (H. sapiens). Body mass estimates for Homo footprint samples appear reasonable when compared to skeletal estimates for related samples. However, estimates for the Laetoli footprint sample using the new formulae appear to be too high when compared to skeletal estimates for A. afarensis. Based on the proportions of A.L. 288-1, this is apparently a result of relatively large feet in this taxon. A different method using a ratio between body mass and foot area in A.L. 288-1 provides estimates more concordant with skeletal estimates and should be used for A. afarensis.

Making the case for possible hominin footprints from the Late Miocene (c. 5.7 Ma) of Crete?

Making the case for possible hominin footprints from the Late Miocene (c. 5.7 Ma) of Crete?