Response to Comment on "Evidence of humans in North America during the Last Glacial Maximum".

Madsen et al. question the reliability of calibrated radiocarbon ages associated with human footprints discovered recently in White Sands National Park, New Mexico, USA. On the basis of the geologic, hydrologic, stratigraphic, and chronologic evidence, we maintain that the ages are robust and conclude that the footprints date to between ~23,000 and 21,000 years ago.Madsen et al. (1) question the veracity of calibrated radiocarbon ages used to constrain the antiquity of human trackways discovered recently at White Sands National Park (WHSA) Locality 2, New Mexico, USA (2). The ages were derived from seeds of the aquatic plant Ruppia cirrhosa, which they suggest may suffer from hard-water (or reservoir) effects, making them too old, potentially by thousands of years. We were well aware of this possibility, investigated it, and presented several lines of evidence that argued against such a problem. Here we respond to each of their four primary points.

Evidence of humans in North America during the Last Glacial Maximum.

Archaeologists and researchers in allied fields have long sought to understand human colonization of North America. Questions remain about when and how people migrated, where they originated, and how their arrival affected the established fauna and landscape. Here, we present evidence from excavated surfaces in White Sands National Park (New Mexico, United States), where multiple in situ human footprints are stratigraphically constrained and bracketed by seed layers that yield calibrated radiocarbon ages between ~23 and 21 thousand years ago. These findings confirm the presence of humans in North America during the Last Glacial Maximum, adding evidence to the antiquity of human colonization of the Americas and providing a temporal range extension for the coexistence of early inhabitants and Pleistocene megafauna.

Sexing white 2D footprints using convolutional neural networks.

Footprints are left, or obtained, in a variety of scenarios from crime scenes to anthropological investigations. Determining the sex of a footprint can be useful in screening such impressions and attempts have been made to do so using single or multi landmark distances, shape analyses and via the density of friction ridges. Here we explore the relative importance of different components in sexing two-dimensional foot impressions namely, size, shape and texture. We use a machine learning approach and compare this to more traditional methods of discrimination. Two datasets are used, a pilot data set collected from students at Bournemouth University (N = 196) and a larger data set collected by podiatrists at Sheffield NHS Teaching Hospital (N = 2677). Our convolutional neural network can sex a footprint with accuracy of around 90% on a test set of N = 267 footprint images using all image components, which is better than an expert can achieve. However, the quality of the impressions impacts on this success rate, but the results are promising and in time it may be possible to create an automated screening algorithm in which practitioners of whatever sort (medical or forensic) can obtain a first order sexing of a two-dimensional footprint.

Earliest parietal art: hominin hand and foot traces from the middle Pleistocene of Tibet.

At Quesang on the Tibetan Plateau we report a series of hand and foot impressions that appear to have been intentionally placed on the surface of a unit of soft travertine. The travertine was deposited by water from a hot spring which is now inactive and as the travertine lithified it preserved the traces. On the basis of the sizes of the hand and foot traces, we suggest that two track-makers were involved and were likely children. We interpret this event as a deliberate artistic act that created a work of parietal art. The travertine unit on which the traces were imprinted dates to between ∼169 and 226 ka BP. This would make the site the earliest currently known example of parietal art in the world and would also provide the earliest evidence discovered to date for hominins on the High Tibetan Plateau (above 4000 m a.s.l.). This remarkable discovery adds to the body of research that identifies children as some of the earliest artists within the genus Homo.

3-D radar imaging unlocks the untapped behavioral and biomechanical archive of Pleistocene ghost tracks.

Footprint evidence of human-megafauna interactions remains extremely rare in the archaeological and palaeontological records. Recent work suggests ancient playa environments may hold such evidence, though the prints may not be visible. These so-called "ghost tracks" comprise a rich archive of biomechanical and behavioral data that remains mostly unexplored. Here we present evidence for the successful detection and 3-D imaging of such footprints via ground-penetrating radar (GPR), including co-associated mammoth and human prints. Using GPR we have found that track density and faunal diversity may be much greater than realized by the unaided human eye. Our data further suggests that detectable subsurface consolidation below mammoth tracks correlates with typical plantar pressure patterns from extant elephants. This opens future potential for more sophisticated biomechanical studies on the footprints of other extinct land vertebrates. Our approach allows rapid detection and documentation of footprints while enhancing the data available from these fossil archives.

Footprints preserve terminal Pleistocene hunt? Human-sloth interactions in North America.

Predator-prey interactions revealed by vertebrate trace fossils are extremely rare. We present footprint evidence from White Sands National Monument in New Mexico for the association of sloth and human trackways. Geologically, the sloth and human trackways were made contemporaneously, and the sloth trackways show evidence of evasion and defensive behavior when associated with human tracks. Behavioral inferences from these trackways indicate prey selection and suggest that humans were harassing, stalking, and/or hunting the now-extinct giant ground sloth in the terminal Pleistocene.

Archaeology and ichnology at Gombore II-2, Melka Kunture, Ethiopia: everyday life of a mixed-age hominin group 700,000 years ago.

We report the occurrence at 0.7 million years (Ma) of an ichnological assemblage at Gombore II-2, which is one of several archaeological sites at Melka Kunture in the upper Awash Valley of Ethiopia, 2000 m asl. Adults and children potentially as young as 12 months old left tracks in a silty substrate on the shore of a body of water where ungulates, as well as other mammals and birds, congregated. Furthermore, the same layers contain a rich archaeological and palaeontological record, confirming that knapping was taking place in situ and that stone tools were used for butchering hippo carcasses at the site. The site gives direct information on hominin landscape use at 0.7 Ma and may provide fresh perspective on the childhood of our ancestors.

Stat-tracks and mediotypes: powerful tools for modern ichnology based on 3D models.

Vertebrate tracks are subject to a wide distribution of morphological types. A single trackmaker may be associated with a range of tracks reflecting individual pedal anatomy and behavioural kinematics mediated through substrate properties which may vary both in space and time. Accordingly, the same trackmaker can leave substantially different morphotypes something which must be considered in creating ichnotaxa. In modern practice this is often captured by the collection of a series of 3D track models. We introduce two concepts to help integrate these 3D models into ichnological analysis procedures. The mediotype is based on the idea of using statistically-generated three-dimensional track models (median or mean) of the type specimens to create a composite track to support formal recognition of a ichno type. A representative track (mean and/or median) is created from a set of individual reference tracks or from multiple examples from one or more trackways. In contrast, stat-tracks refer to other digitally generated tracks which may explore variance. For example, they are useful in: understanding the preservation variability of a given track sample; identifying characteristics or unusual track features; or simply as a quantitative comparison tool. Both concepts assist in making ichnotaxonomical interpretations and we argue that they should become part of the standard procedure when instituting new ichnotaxa. As three-dimensional models start to become a standard in publications on vertebrate ichnology, the mediotype and stat-track concepts have the potential to help guiding a revolution in the study of vertebrate ichnology and ichnotaxonomy.

Animal movements in the Kenya Rift and evidence for the earliest ambush hunting by hominins.

Animal movements in the Kenya Rift Valley today are influenced by a combination of topography and trace nutrient distribution. These patterns would have been the same in the past when hominins inhabited the area. We use this approach to create a landscape reconstruction of Olorgesailie, a key site in the East African Rift with abundant evidence of large-mammal butchery between ~1.2 and ~0.5 Ma BP. The site location in relation to limited animal routes through the area show that hominins were aware of animal movements and used the location for ambush hunting during the Lower to Middle Pleistocene. These features explain the importance of Olorgesailie as a preferred location of repeated hominin activity through multiple changes in climate and local environmental conditions, and provide insights into the cognitive and hunting abilities of Homo erectus while indicating that their activities at the site were aimed at hunting, rather than scavenging.

Landscapes and their relation to hominin habitats: case studies from Australopithecus sites in eastern and southern Africa.

We examine the links between geomorphological processes, specific landscape features, surface water drainage, and the creation of suitable habitats for hominins. The existence of mosaic (i.e., heterogeneous) habitats within hominin site landscape reconstructions is typically explained using models of the riverine and gallery forest settings, or the pan or lake setting. We propose a different model: the Tectonic Landscape Model (TLM), where tectonic faulting and volcanism disrupts existing pan or river settings at small-scales (∼10-25 km). Our model encompasses the interpretation of the landscape features, the role of tectonics in creating these landscapes, and the implications for hominins. In particular, the model explains the underlying mechanism for the creation and maintenance of heterogeneous habitats in regions of active tectonics. We illustrate how areas with faulting and disturbed drainage patterns would have been attractive habitats for hominins, such as Australopithecus, and other fauna. Wetland areas are an important characteristic of surface water disturbance by fault activity; therefore we examine the tectonically-controlled Okavango Delta (Botswana) and the Nylsvley wetland (South Africa) as modern examples of how tectonics in a riverine setting significantly enhance the faunal and floral biodiversity. While tectonic landscapes may not have been the only type of attractive habitats to hominins, we propose a suite of landscape, faunal, and floral indicators, which when recovered together suggest that site environments may have been influenced by tectonic and/or volcanic activity while hominins were present. For the fossil sites, we interpret the faulting and landscapes around australopithecine-bearing sites of the Middle Awash (Ethiopia) and Makapansgat, Taung, and Sterkfontein (South Africa) to illustrate these relationships between landscape features and surface water bodies. Exploitation of tectonically active landscapes may explain why the paleoenvironmental signals, anatomy, diets, as well as the fauna associated with Australopithecus appear largely heterogeneous through time and space. This hypothesis is discussed in light of potential preservation and time-averaging effects which may affect patterns visible in the fossil record. The model, however, offers insight into the landscape processes of how such habitats are formed. The landscape features and range of habitat conditions, specifically the wetter, down-dropped plains and drier, uplifted flanks persist in close proximity for as long as the fault motion continues. The Tectonic Landscape Model provides an alternative explanation of why mixed habitats may be represented at certain sites over longer timescales.

Landscapes of human evolution: models and methods of tectonic geomorphology and the reconstruction of hominin landscapes.

This paper examines the relationship between complex and tectonically active landscapes and patterns of human evolution. We show how active tectonics can produce dynamic landscapes with geomorphological and topographic features that may be critical to long-term patterns of hominin land use, but which are not typically addressed in landscape reconstructions based on existing geological and paleoenvironmental principles. We describe methods of representing topography at a range of scales using measures of roughness based on digital elevation data, and combine the resulting maps with satellite imagery and ground observations to reconstruct features of the wider landscape as they existed at the time of hominin occupation and activity. We apply these methods to sites in South Africa, where relatively stable topography facilitates reconstruction. We demonstrate the presence of previously unrecognized tectonic effects and their implications for the interpretation of hominin habitats and land use. In parts of the East African Rift, reconstruction is more difficult because of dramatic changes since the time of hominin occupation, while fossils are often found in places where activity has now almost ceased. However, we show that original, dynamic landscape features can be assessed by analogy with parts of the Rift that are currently active and indicate how this approach can complement other sources of information to add new insights and pose new questions for future investigation of hominin land use and habitats.