The Laetoli footprints and early hominin locomotor kinematics

A critical question in human evolution is whether the earliest bipeds walked with a bent-hip, bent-knee gait or on more extended hindlimbs. The differences between these gaits are not trivial, because the adoption of either has important implications for the evolution of bipedalism. In this study, we re-examined the Laetoli footprints to determine whether they can provide information on the locomotor posture of early hominins. Previous researchers have suggested that the stride lengths of Laetoli hominins fall within the range of modern human stride lengths and therefore, Laetoli hominins walked with modern-human-like kinematics. Using a dynamic-similarity analysis, we compared Laetoli hominin stride lengths with those of both modern humans and chimpanzees. Our results indicate that Laetoli hominins could have used either a bent-hip, bent-knee gait, similar to a chimpanzee, or an extended-hindlimb gait, similar to a human. In fact, our data suggest that the Laetoli hominins could have walked near their preferred speeds using either limb posture. This result contrasts with most previous studies, which suggest relatively slow walking speeds for these early bipeds. Despite the many attempts to discern limb-joint kinematics from Laetoli stride lengths, our study concludes that stride lengths alone do not resolve the debate over early hominin locomotor postures.     

The semicircular canal system and locomotion: The case of extinct lemuroids and lorisoids

Paleontologists reconstruct the locomotor and postural behavior of extinct species by analogy with living forms and biomechanical analyses. In rare cases, behavioral evidence such as footprints can be used to confirm fossil‐based reconstructions for predominantly terrestrial orders of mammals. For instance, the chalicothere prints from Laetoli show that these perissodactyls supported their body weight on the metacarpals, as previously reconstructed. 1 Unfortunately, primates are mostly arboreal and rarely leave footprints. The cercopithecid and hominin prints at Laetoli are a rare exception. We have recently shown that the semicircular canal system can be used to test and augment locomotor reconstructions based on postcranial material or to provide first estimations of locomotor behavior for taxa not known from the postcranium. Using a sample of modern primates, we have been able to demonstrate that the radii of curvature of the semicircular canals are significantly correlated with both body mass and agility of locomotion. 2 This paper reviews those results and examines the relationship between semicircular canal morphology and other evidence in efforts to reconstruct locomotor behavior in subfossil lemurs from the Holocene of Madagascar and fossil lorisoids from the Miocene of Africa.     

Hominid footprints at Laetoli: facts and interpretations

The history of discovery and interpretation of primate footprints at the site of Laetoli in northern Tanzania is reviewed. An analysis of the geological context of these tracks is provided. The hominid tracks in Tuff 7 at Site G in the Garusi River Valley demonstrate bipedality at a mid-Pliocene datum. Comparison of these tracks and the Hadar hominid foot fossils by Tuttle has led him to conclude that Australopithecus afarensis did not make the Tanzanian prints and that a more derived form of hominid is therefore indicated at Laetoli. An alternative interpretation has been offered by Stern and Susman who posit a conforming "transitional morphology" in both the Tanzanian prints and the Ethiopian bones. The present examines both hypotheses and shows that neither is likely to be entirely correct. To illustrate this point, a reconstruction of the foot skeleton of a female A. afarensis is undertaken, and the results are compared to the Laetoli tracks. We conclude that A. afarensis represents the best candidate for the maker of the Laetoli hominid trails.     

Stride length and its determinants in humans, early hominids, primates, and mammals

Primate stride lengths during quadrupedal locomotion are very long when compared to those of nonprimate quadrupedal mammals at the speed of trot/gallop transition. These exceptional lengths are a consequence of the relatively long limbs of primates and the large angular excursions of their limbs during quadrupedalism. When quadrupedal primates employ bipedal gaits they exhibit much lower angular excursions. Consequently their bipedal stride lengths do not appear to be exceptional in length when compared to other mammals. Angular excursions of the lower limbs of modern humans are not exceptionally large. However, when running, humans exhibit relatively long periods of flight (i.e., they have low duty factors) when compared to other mammals including primates. Because of these long periods of flight and their relative long lower limbs, humans have running stride lengths that are at the lower end of the range of stride lengths of quadrupedal primates. The stride length of the Laetoli hominid trails are evaluated in this context.     

Creeping patterns of human adults and infants

The patterns of swing and support for the hands-and-feet or hands-and-knees gaits (creeping) of human adults and infants are compared based on data from a number of studies. Human infants show considerable variability in their creeping gait patterns, whereas adult patterns are less variable and fairly consistent after a few minutes of practice. Creeping on hands-and-knees appears to dictate a gait pattern characteristically different from creeping on hands-and-feet. The highly inefficient nature of adult creeping supports the view that our early hominid ancestors were poorly adapted to quadrupedal locomotion. Data obtained from high-speed film analysis of human creeping patterns show that the number of foot lengths per stride in creeping is about twice that for normal upright walking at the same speed. The support pattern of human creeping is different from that of nonhuman primates. These findings are discussed in the context of debate concerning the origin of the Laetoli hominid footprints and the knuckle-walking hypothesis.     

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.     

History of Ichnology: The Misconceived Footprints of Rhynchosaurs

In 1842 Richard Owen described a Triassic reptile from Grinshill, Shropshire, which he named Rhynchosaurus articeps. He suggested that footprints found in the same beds were those of this fossil. However, the footprints were characterised by a backward-pointing toe and so were of the type now known as Rotodactylus. Huxley (1877), Woodward (1907), and Benton (1990) have subsequently shown that the five digits of Rhynchosaurus point forward and so could not have left these footprints. In 1896 Beasley classified the Triassic footprints found in Cheshire, his type D prints being those earlier assigned to rhynchosaurs. His D1 prints were named Rhynchosauroides articeps by Maidwell (1911). However, these D1 prints, which come from a lower horizon in the Anisian, are consistently too small to match Owen's fossil. Beasley's D3 form, now named Synaptichnium pseudosuchoides Nopcsa, is more likely to represent the footprints of Rhynchosaurus articeps, although further research and study of more complete trackways will be necessary to clarify whether these are the footprints of Archosauromorphs, such as rhynchosaurs or possibly those of Archosauriformes, for example, erythrosuchids or proterosuchids. Maidwell's Rhynchosauroides rectipes and Rhynchosauroides membranipes, originally believed to be distinct ichnospecies, are more likely to be synonyms, their apparent differences reflecting variations in the substrate traversed.     

Pliocene Animal Trackways at Laetoli: Research and Conservation Potential

Laetoli, a paleoanthropological site in Northern Tanzania, is perhaps best known for its famous fossil hominid footprints that were discovered by Mary Leakey and her co-workers in 1978. The site not only preserves the hominid footprints but also trackways, which provide a snapshot of Pliocene faunal communities from East Africa and their inferred environments. Unlike the hominid footprints at site G, which have received tremendous attention, the animal trackways, especially at Localities 7, 8 and 10 have been neglected and are fast disappearing. In this paper, we discuss animal tracks at a newly discovered exposure and provide preliminary data on the tracks at this exposure and other sites. We also discuss the importance of the animal trackways as ecological indicators, which we have investigated as part of ongoing research and conservation efforts initiated by the Tanzania Field School in Paleoanthropology and the Associated Colleges of the Midwest (ACM) Tanzania Semester Abroad programs.     

The locomotor anatomy of Australopithecus afarensis

The postcranial skeleton of Australopithecus afarensis from the Hadar Formation, Ethiopia, and the footprints from the Laetoli Beds of northern Tanzania, are analyzed with the goal of determining (1) the extent to which this ancient hominid practiced forms of locomotion other than terrestrial bipedality, and (2) whether or not the terrestrial bipedalism of A. afarensis was notably different from that of modern humans. It is demonstrated that A. afarensis possessed anatomic characteristics that indicate a significant adaptation for movement in the trees. Other structural features point to a mode of terrestrial bipedality that involved less extension at the hip and knee than occurs in modern humans, and only limited transfer of weight onto the medial part of the ball of the foot, but such conclusions remain more tentative than that asserting substantive arboreality. A comparison of the specimens representing smaller individuals, presumably female, to those of larger individuals, presumably male, suggests sexual differences in locomotor behavior linked to marked size dimorphism. The males were probably less arboreal and engaged more frequently in terrestrial bipedalism. In our opinion, A. afarensis from Hadar is very close to what can be called a "missing link." We speculate that earlier representatives of the A. afarensis lineage will present not a combination of arboreal and bipedal traits, but rather the anatomy of a generalized ape.     

An enigmatic early mesozoic dinosaur trackway from Zimbabwe

A trackway from Zimbabwe of probably the smallest dinosaur footprints recorded in Africa, is described and tentatively assigned to the Early Jurassic. The footprints are possibly those of a theropod and show strong negative (outward) rotation of the pes and are associated with manus prints. The shape of the footprints, unusual negative rotation, posterior curvature of digit IV and curious positioning of the manus prints in relation to the pes are enigmatic but somewhat reminiscent of Atreipus. Although a number of propositions are considered the most likely is that the animal was an immature dinosaur using a quadrupedal gait. A second trackway of slightly larger footprints of a bipedal theropod dinosaur is also recorded along with other diminutive tracks that suggest an early dinosaur assemblage, possibly dating from near the Trias‐sic‐Jurassic boundary.     

Laetoli toes andAustralopithecus afarensis

The probable misfit between feet, particularly toes II–V, of 3.0-million-year-oldAustralopithecus afarensis from Hadar, Ethiopia, and the 3.5-million-year-old hominid footprints at Site G, Laetoli, Tanzania, casts doubt thatA. Afarensis made the Laetoli trails. We suggest that another species ofAustralopithecus or an anonymous genus of the Hominidae, with remarkably humanoid feet, walked at Laetoli. It would be imprudent to declare thatHomo was present at Laetoli 3.5 million years ago (my) because there is no evidence of brain expansion, advanced tool manufacture, or other non-locomotor hallmarks of the human condition at Site G.     

Pleistocene human footprints from the Willandra Lakes, Southeastern Australia

Human and other hominid fossil footprints provide rare but important insights into anatomy and behavior. Here we report recently discovered fossil trackways of human footprints from the Willandra Lakes region of western New South Wales, Australia. Optically dated to between 19-23 ka and consisting of at least 124 prints, the trackways form the largest collection of Pleistocene human footprints in the world. The prints were made by adults, adolescents, and children traversing the moist surface of an ephemeral soak. This site offers a unique glimpse of humans living in the arid inland of Australia at the height of the last glacial period.     

Using Patterns in Track-Plate Footprints to Identify Individual Fishers

Abstract: If individuals can be identified from patterns in their footprints, noninvasive survey methods can be used to estimate abundance. Track plates capture fine detail in the footprints of fishers (Martes pennanti), recording rows of dots corresponding to tiny papillae on the animal's metacarpal pad. We show that the pattern of these dots can be used to identify individual fishers, similar to human fingerprints. A probabilistic model of uniqueness based on variation in spacing between 1,400 pairs of dots that we measured in prints of 14 different fisher feet suggests the probability of encountering a similar pattern in the print of a different foot by chance alone is ≤ 0.35ⁿ, where n = the number of dot pairs examined. This predicts a 0.00003 probability that a match made using 10 pairs of dots is false. Dot spacing from footprints made by the same foot was remarkably consistent (sN = 0.02 mm, n = 24 dot pairs). Combined, these results suggest dot patterns in fisher footprints were unique to individuals and were consistently reproduced on track plates. Empirical tests of matching accuracy were best with good-quality prints, highlighting the need for experience judging when prints are usable. We applied print matching to fisher detections collected on track plates deployed at 500-m intervals along 10 3.5-km transects in the Adirondack region of New York, USA. Of 62 fisher detections, 85% had ≥ 1 footprint of suitable quality to compare with other high-quality prints. We found that most detections from a transect were from the same individual fisher suggesting nonindependence of detections. Thus, data from traditional track-plate deployments over small time periods cannot be used as a measure of abundance, but new study designs using print matching could obtain robust noninvasive, mark—recapture density estimates.     

Impact of sauropod dinosaurs on lagoonal substrates in the Broome Sandstone (Lower Cretaceous), Western Australia

Existing knowledge of the tracks left by sauropod dinosaurs (loosely 'brontosaurs') is essentially two-dimensional, derived mainly from footprints exposed on bedding planes, but examples in the Broome Sandstone (Early Cretaceous) of Western Australia provide a complementary three-dimensional picture showing the extent to which walking sauropods could deform the ground beneath their feet. The patterns of deformation created by sauropods traversing thinly-stratified lagoonal deposits of the Broome Sandstone are unprecedented in their extent and structural complexity. The stacks of transmitted reliefs (underprints or ghost prints) beneath individual footfalls are nested into a hierarchy of deeper and more inclusive basins and troughs which eventually attain the size of minor tectonic features. Ultimately the sauropod track-makers deformed the substrate to such an extent that they remodelled the topography of the landscape they inhabited. Such patterns of substrate deformation are revealed by investigating fragmentary and eroded footprints, not by the conventional search for pristine footprints on intact bedding planes. For that reason it is not known whether similar patterns of substrate deformation might occur at sauropod track-sites elsewhere in the world.     

Trackways of the American Crocodile (Crocodylus acutus) in Northwestern Costa Rica: Implications for Crocodylian Ichnology

We documented trackways of free-living Crocodylus acutus on beaches at the mouths of Tamarindo and Ventanas estuaries, Costa Rica. Our crocodiles had estimated total lengths of 1–3 meters or more. Manus prints have five digits, with digits I–III bearing claw marks. Pes prints have four digits, with claw marks on digits I–III. The pes is plantigrade. Claws generally dig into the substrate. Apart from claw marks, digit I and the heel of the pes are usually the most deeply impressed parts of footprints. Trackways are wide-gauge. Pes prints are usually positioned just behind ipsilateral manus prints of the same set and may overlap them. Manus and pes prints angle slightly outward with respect to the crocodile's direction of movement. Claw-bearing digits of both the manus and pes may create curved, concave-toward-the-midline drag marks as the autopodium is protracted. The tail mark varies in depth and clarity, and in shape from nearly linear to markedly sinuous. Sometimes the tail mark hugs the trackway midline, but sometimes it is closer to, or even cuts across, prints of one side. American crocodile footprints and trackways are similar to those observed in other extant crocodylian species, indicating substantial trackway conservatism across the group.     

A study on continuous follow-the-leader (FTL) gaits: an effective walking algorithm over rough terrain

The follow-the-leader (FTL) gait is an effective walking algorithm for a legged system to traverse a rough terrain. In an FTL gait, all the legs simply place at the footprints made by the legs ahead of them. By this way the demand on foothold selection is significantly reduced. A special category of FTL gaits, called continuous FTL gaits, provide a smooth body motion during walking and enable the legged system to reach a higher speed. In this paper, a comprehensive study of continuous FTL gaits is presented. The equations for two types of continuous FTL gaits are formulated. The stability of these continuous FTL gaits is studied analytically and verified numerically. Strategies of forbidden area avoidance and special methods of large foot adjustment are introduced. The motion resulting from the use of these strategies and methods is simulated and checked using computer graphics.     

Natural gaits of the non-pathological flat foot and high-arched foot

There has been a controversy as to whether or not the non-pathological flat foot and high-arched foot have an effect on human walking activities. The 3D foot scanning system was employed to obtain static footprints from subjects adopting a half-weight-bearing stance. Based upon their footprints, the subjects were divided into two groups: the flat-footed and the high-arched. The plantar pressure measurement system was used to measure and record the subjects' successive natural gaits. Two indices were proposed: distribution of vertical ground reaction force (VGRF) of plantar and the rate of change of footprint areas. Using these two indices to compare the natural gaits of the two subject groups, we found that (1) in stance phase, there is a significant difference (p<0.01) in the distributions of VGRF of plantar; (2) in a stride cycle, there is also a significant difference (p<0.01) in the rate of change of footprint area. Our analysis suggests that when walking, the VGRF of the plantar brings greater muscle tension to the flat-footed while a smaller rate of change of footprint area brings greater stability to the high-arched.     

Modeling three-dimensional sculptures of australopithecines (Australopithecus afarensis) for the Museum of Natural History of Vienna (Austria): the post-cranial hypothesis.

In March 1999, E. Daynes, a sculptor specializing in fossil hominid reconstruction, asked C. Berge to take over the scientific supervision of the reconstruction of two australopithecine post-crania. The heads had been modeled from two skulls found in Hadar (AL 444-2, AL 417). The sculptures were to be represented in a walking stance. The female proportions (AL 417) are estimated from the skeleton of 'Lucy' (AL 288), and the male proportions (AL 444-2) extrapolated from the female ones. Biomechanical and anatomical data (comparison with great apes and humans) are used to reconstruct both dynamic equilibrium and muscular systems. The reconstruction suggests that the fossils moved the pelvis and shoulders extensively when they walked. The hindlimb muscles (such as adductors, gluteal muscles and calf) are fleshy and not or very little tendinous. As indicated by the Laetoli step prints (belonging to a close and contemporaneous species), the foot is adducted during the walk and the support is internal just before take off. In spite of inevitable approximations, such a reconstruction appears to be particularly helpful to bring out morphological and functional traits of the first hominids which are both close to and different from modern humans.     

Holocene Human Footprints in North America

Reports of human footprints from North American archaeological sites are summarized and described. Although some human footprints have been claimed to be of Pleistocene and earlier ages, the earliest authenticated prints in America north of Mexico, as distinct from Central or South America, date to the Holocene. The earliest of these prints is at least 5,070 years old, and the youngest are late prehistoric period, only about 400 years old. Footprint sites are reported in the U.S. Southeast, Southwest, and California. Activities indicated by ancient North American human footprints include the mundane, such as daily tasks in a riparian zone and domestic behaviors inside a habitation. Footprints were also left by prehistoric peoples undertaking more esoteric activities deep in cave interiors, including exploration, mineral extraction, and ritual.