Testing inter-observer error under a collaborative research framework for studying lithic shape variability.

Evaluating error that arises through the aggregation of data recorded by multiple observers is a key consideration in many metric and geometric morphometric analyses of stone tool shape. One of the most common approaches involves the convergence of observers for repeat trails on the same set of artefacts: however, this is logistically and financially challenging when collaborating internationally and/or at a large scale. We present and evaluate a unique alternative for testing inter-observer error, involving the development of 3D printed copies of a lithic reference collection for distribution among observers. With the aim of reducing error, clear protocols were developed for photographing and measuring the replicas, and inter-observer variability was assessed on the replicas in comparison with a corresponding data set recorded by a single observer. Our results demonstrate that, when the photography procedure is standardized and dimensions are clearly defined, the resulting metric and geometric morphometric data are minimally affected by inter-observer error, supporting this method as an effective solution for assessing error under collaborative research frameworks. Collaboration is becoming increasingly important within archaeological and anthropological sciences in order to increase the accessibility of samples, encourage dual-project development between foreign and local researchers and reduce the carbon footprint of collection-based research. This study offers a promising validation of a collaborative research design whereby researchers remotely work together to produce comparable data capturing lithic shape variability. Supplementary Information: The online version contains supplementary material available at 10.1007/s12520-022-01676-2.

A spatiotemporally explicit paleoenvironmental framework for the Middle Stone Age of eastern Africa.

Eastern Africa has played a prominent role in debates about human evolution and dispersal due to the presence of rich archaeological, palaeoanthropological and palaeoenvironmental records. However, substantial disconnects occur between the spatial and temporal resolutions of these data that complicate their integration. Here, we apply high-resolution climatic simulations of two key parameters, mean annual temperature and precipitation, and a biome model, to produce a highly refined characterisation of the environments inhabited during the eastern African Middle Stone Age. Occupations are typically found in sub-humid climates and landscapes dominated by or including tropical xerophytic shrubland. Marked expansions from these core landscapes include movement into hotter, low-altitude landscapes in Marine Isotope Stage 5 and cooler, high-altitude landscapes in Marine Isotope Stage 3, with the recurrent inhabitation of ecotones between open and forested habitats. Through our use of high-resolution climate models, we demonstrate a significant independent relationship between past precipitation and patterns of Middle Stone Age stone tool production modes overlooked by previous studies. Engagement with these models not only enables spatiotemporally explicit examination of climatic variability across Middle Stone Age occupations in eastern Africa but enables clearer characterisation of the habitats early human populations were adapted to, and how they changed through time.

Evaluating refugia in recent human evolution in Africa.

Homo sapiens have adapted to an incredible diversity of habitats around the globe. This capacity to adapt to different landscapes is clearly expressed within Africa, with Late Pleistocene Homo sapiens populations occupying savannahs, woodlands, coastlines and mountainous terrain. As the only area of the world where Homo sapiens have clearly persisted through multiple glacial-interglacial cycles, Africa is the only continent where classic refugia models can be formulated and tested to examine and describe changing patterns of past distributions and human phylogeographies. The potential role of refugia has frequently been acknowledged in the Late Pleistocene palaeoanthropological literature, yet explicit identification of potential refugia has been limited by the patchy nature of palaeoenvironmental and archaeological records, and the low temporal resolution of climate or ecological models. Here, we apply potential climatic thresholds on human habitation, rooted in ethnographic studies, in combination with high-resolution model datasets for precipitation and biome distributions to identify persistent refugia spanning the Late Pleistocene (130-10 ka). We present two alternate models suggesting that between 27% and 66% of Africa may have provided refugia to Late Pleistocene human populations, and examine variability in precipitation, biome and ecotone distributions within these refugial zones. This article is part of the theme issue 'Tropical forests in the deep human past'.

The Importance of Noise Colour in Simulations of Evolutionary Systems.

Simulations of evolutionary dynamics often employ white noise as a model of stochastic environmental variation. Whilst white noise has the advantages of being simply generated and analytically tractable, empirical analyses demonstrate that most real environmental time series have power spectral densities consistent with pink or red noise, in which lower frequencies contribute proportionally greater amplitudes than higher frequencies. Simulated white noise environments may therefore fail to capture key components of real environmental time series, leading to erroneous results. To explore the effects of different noise colours on evolving populations, a simple evolutionary model of the interaction between life-history and the specialism-generalism axis was developed. Simulations were conducted using a range of noise colours as the environments to which agents adapted. Results demonstrate complex interactions between noise colour, reproductive rate, and the degree of evolved generalism; importantly, contradictory conclusions arise from simulations using white as opposed to red noise, suggesting that noise colour plays a fundamental role in generating adaptive responses. These results are discussed in the context of previous research on evolutionary responses to fluctuating environments, and it is suggested that Artificial Life as a field should embrace a wider spectrum of coloured noise models to ensure that results are truly representative of environmental and evolutionary dynamics.

Neural networks differentiate between Middle and Later Stone Age lithic assemblages in eastern Africa.

The Middle to Later Stone Age transition marks a major change in how Late Pleistocene African populations produced and used stone tool kits, but is manifest in various ways, places and times across the continent. Alongside changing patterns of raw material use and decreasing artefact sizes, changes in artefact types are commonly employed to differentiate Middle Stone Age (MSA) and Later Stone Age (LSA) assemblages. The current paper employs a quantitative analytical framework based upon the use of neural networks to examine changing constellations of technologies between MSA and LSA assemblages from eastern Africa. Network ensembles were trained to differentiate LSA assemblages from Marine Isotope Stage 3&4 MSA and Marine Isotope Stage 5 MSA assemblages based upon the presence or absence of 16 technologies. Simulations were used to extract significant indicator and contra-indicator technologies for each assemblage class. The trained network ensembles classified over 94% of assemblages correctly, and identified 7 key technologies that significantly distinguish between assemblage classes. These results clarify both temporal changes within the MSA and differences between MSA and LSA assemblages in eastern Africa.

Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?

We challenge the view that our species, Homo sapiens, evolved within a single population and/or region of Africa. The chronology and physical diversity of Pleistocene human fossils suggest that morphologically varied populations pertaining to the H. sapiens clade lived throughout Africa. Similarly, the African archaeological record demonstrates the polycentric origin and persistence of regionally distinct Pleistocene material culture in a variety of paleoecological settings. Genetic studies also indicate that present-day population structure within Africa extends to deep times, paralleling a paleoenvironmental record of shifting and fractured habitable zones. We argue that these fields support an emerging view of a highly structured African prehistory that should be considered in human evolutionary inferences, prompting new interpretations, questions, and interdisciplinary research directions.

Climatic variability, plasticity, and dispersal: A case study from Lake Tana, Ethiopia.

The numerous dispersal events that have occurred during the prehistory of hominin lineages are the subject of longstanding and increasingly active debate in evolutionary anthropology. As well as research into the dating and geographic extent of such dispersals, there is an increasing focus on the factors that may have been responsible for dispersal. The growing body of detailed regional palaeoclimatic data is invaluable in demonstrating the often close relationship between changes in prehistoric environments and the movements of hominin populations. The scenarios constructed from such data are often overly simplistic, however, concentrating on the dynamics of cyclical contraction and expansion during severe and ameliorated conditions respectively. This contribution proposes a two-stage hypothesis of hominin dispersal in which populations (1) accumulate high levels of climatic tolerance during highly variable climatic phases, and (2) express such heightened tolerance via dispersal in subsequent low-variability phases. Likely dispersal phases are thus proposed to occur during stable climatic phases that immediately follow phases of high climatic variability. Employing high resolution palaeoclimatic data from Lake Tana, Ethiopia, the hypothesis is examined in relation to the early dispersal of Homo sapiens out of East Africa and into the Levant. A dispersal phase is identified in the Lake Tana record between c. 112,550 and c. 96,975 years ago, a date bracket that accords well with the dating evidence for H. sapiens occupation at the sites of Qafzeh and Skhul. Results are discussed in relation to the complex pattern of H. sapiens dispersal out of East Africa, with particular attention paid to the implications of recent genetic chronologies for the origin of non-African modern humans.

Recurrent patterning in the daily foraging routes of hamadryas baboons (Papio hamadryas): spatial memory in large-scale versus small-scale space.

The benefits of spatial memory for foraging animals can be assessed on two distinct spatial scales: small-scale space (travel within patches) and large-scale space (travel between patches). While the patches themselves may be distributed at low density, within patches resources are likely densely distributed. We propose, therefore, that spatial memory for recalling the particular locations of previously visited feeding sites will be more advantageous during between-patch movement, where it may reduce the distances traveled by animals that possess this ability compared to those that must rely on random search. We address this hypothesis by employing descriptive statistics and spectral analyses to characterize the daily foraging routes of a band of wild hamadryas baboons in Filoha, Ethiopia. The baboons slept on two main cliffs--the Filoha cliff and the Wasaro cliff--and daily travel began and ended on a cliff; thus four daily travel routes exist: Filoha-Filoha, Filoha-Wasaro, Wasaro-Wasaro, Wasaro-Filoha. We use newly developed partial sum methods and distribution-fitting analyses to distinguish periods of area-restricted search from more extensive movements. The results indicate a single peak in travel activity in the Filoha-Filoha and Wasaro-Filoha routes, three peaks of travel activity in the Filoha-Wasaro routes, and two peaks in the Wasaro-Wasaro routes; and are consistent with on-the-ground observations of foraging and ranging behavior of the baboons. In each of the four daily travel routes the "tipping points" identified by the partial sum analyses indicate transitions between travel in small- versus large-scale space. The correspondence between the quantitative analyses and the field observations suggest great utility for using these types of analyses to examine primate travel patterns and especially in distinguishing between movement in small versus large-scale space. Only the distribution-fitting analyses are inconsistent with the field observations, which may be due to the scale at which these analyses were conducted.

The evolution of spatial memory.

Models of foraging behaviour often assume either that animals are searching for resources, and therefore have no prior knowledge of resource locations, or that they are effectively omniscient, with a comprehensive knowledge of their habitat. By contrast, few attempts have been made to examine the actual conditions under which spatial memory will provide net benefits to foragers. To redress this balance, a model is developed that relates the sensory acuity of the forager and key indices of resource structure to the expected foraging efficiency via calculation of inter-patch distances. Efficiencies of 'ignorant' and 'prescient' foragers are examined in order to derive sets of conditions under which natural selection will favour the evolution of spatial memory capabilities. Results suggest that when resources are densely distributed or sensory acuity is high, spatial memory for resource locations provides no increase in efficiency over that of an 'ignorant' forager encountering resources at random.

Food and social complexity at Çayönü Tepesi, southeastern Anatolia: Stable isotope evidence of differentiation in diet according to burial practice and sex in the early Neolithic.

The identification of early social complexity and differentiation in early village societies has been approached in the past most notably through the evaluation of rituals and architectural layouts. Such studies could be complemented by an approach that provides data about everyday behaviours of individuals. We took 540 human and animal bone samples for stable carbon and nitrogen isotope analysis from the Neolithic site of Çayönü Tepesi in southeastern Anatolia. The inhabitants at this site chose to bury their dead in two different ways at different times during its occupation: beneath the floors of their houses, but also inside a public mortuary building known as the Skull Building. This variation provides an opportunity using isotope methods to test whether there was evidence for structuring of daily activities (diet in this case) that might serve to reinforce this change in burial practice. We show that when the inhabitants of Çayönü Tepesi changed their architecture and operated different burial practices in conjunction, this coincided with other aspects of behaviour including socially-constituted food consumption practices, which served to reinforce social identities.

Fission-fusion and the evolution of hominin social systems.

The course of hominin evolution has involved successive migrations towards higher absolute latitudes over the past three million years. Poorer habitat quality further from the equator has led to the necessity for groups occupying higher latitudes to live at lower population densities. Coupled with a trend towards increasing group size over this time period, this tendency towards expansion has led to exponential increases in the area requirements of hominin groups, and a concomitant need to adjust foraging patterns. The current analyses suggest that the development of increasingly complex, multi-level fission-fusion social systems could have freed hominins of the foraging constraints imposed by large group sizes and low population densities. Analyses of the fossil record suggest latitudinally-driven differences in area requirements of the australopithecines from East and South Africa, and African and Asian Homo erectus. In contrast, chronologically-driven differences appear between H. erectus as a whole and Homo heidelbergensis, and between H. heidelbergensis and the Neanderthals. These results are discussed in relation to studies of the foraging patterns of primates and hunter-gatherers.

Speciation, diversity, and Mode 1 technologies: the impact of variability selection.

Over geological timescales, organisms encounter periodic shifts in selective conditions driven by environmental change. The variability selection hypothesis suggests that increases in environmental fluctuation have led to the evolution of complex, flexible behaviours able to respond to novel and unpredictable adaptive settings. This hypothesis is tested via the framework of a single locus genetic model in which an invading 'versatilist' allele competes with two opposed specialists in a selection regime driven by a fluctuating environment, modelled initially as a sine wave and subsequently as an empirical climate curve covering the past 5 million years. Results demonstrate that generalist alleles achieve fixation in the sine wave environment, whilst versatilist alleles do so in the empirical environment, even at a range of very low fitness advantages over the basic generalist template. Variability selection is found to be a particularly strong force between approximately 2.5 and 1.2 Ma (millions of years ago). These results are discussed in relation to the spread of Oldowan lithics and the patterns of speciation and extinction documented in the hominin fossil record. It is suggested that the flexibility required for survival in a variable climatic regime may have been a stimulus to the development of the first stone tool technologies, whilst the ecological opportunities provided by heightened variability may have been a factor in prompting the hominin adaptive radiation evidenced during this period.