Late Acheulean occupations at Montagu Cave and the pattern of Middle Pleistocene behavioral change in Western Cape, southern Africa.

Patterns of so-called modern human behavior are increasingly well documented in an abundance of Middle Stone Age archaeological sites across southern Africa. Contextualized archives directly preceding the southern African Middle Stone Age, however, remain scarce. Current understanding of the terminal Acheulean in southern Africa derives from a small number of localities that are predominantly in the central and northern interior. Many of these localities are surface and deflated contexts, others were excavated prior to the availability of modern field documentation techniques, and yet other relevant assemblages contain low numbers of characteristic artifacts relative to volume of excavated deposit. The site of Montagu Cave, situated in the diverse ecosystem of the Cape Floral Region, South Africa, contains the rare combination of archaeologically rich, laminated and deeply stratified Acheulean layers followed by a younger Middle Stone Age occupation. Yet little is known about the site owing largely to a lack of contextual information associated with the early excavations. Here we present renewed excavation of Levels 21-22 at Montagu Cave, located in the basal Acheulean sequence, including new data on site formation and ecological context, geochronology, and technological variability. We document intensive occupation of the cave by Acheulean tool-producing hominins, likely at the onset of interglacial conditions in MIS 7. New excavations at Montagu Cave suggest that, while Middle Stone Age technologies were practiced by 300 ka in several other regions of Africa, the classic Acheulean persisted later in the Fynbos Biome of the southwestern Cape. We discuss the implications of this regionalized persistence for the biogeography of African later Middle Pleistocene hominin populations, for the ecological drivers of their technological systems, and for the pattern and pace of behavioral change just prior to the proliferation of the southern African later Middle Stone Age.

Quantifying differences in hominin flaking technologies with 3D shape analysis.

Genetic and climate-driven estimates of past population dynamics are increasingly influential in broader models of hominin migration and adaptation, yet the contribution of stone artifact variability remains more contentious. Scientists are increasingly recognizing the potential of unretouched stone flakes ('flakes') in exploring existing models of hominin behavioral evolution. This is because flakes (1) were produced by all stone tool manufacturing groups in the past, (2) are abundant from the inception of the archaeological record up into the ethnographic present, and (3) preserve under most conditions. The statistical tools of 3D geometric morphometrics capture detailed approximations of flake form that are challenging to document with conventional artifact analyses. We analyze a collection of 717 3D scans of experimentally produced flakes from 5 production strategies that were practiced by hominins through large parts of the Pleistocene and that scientists have drawn on also to make demographic arguments about past human behavior (n = 45 reduction sequences, n = 3 knappers naive toward the study objectives). First, as a proof of concept, we demonstrate that we can estimate the strategies used to produce these flakes at a high success rate even when flakes from early stages of core reduction are included. We frame the significance of this finding against archaeological classifications from several key Middle Paleolithic assemblages in France (n = 4 sites, n = 28 layers, n = 16,467 flakes). Second, we show that 3D geometric morphometrics captures subtle differences in these strategies that influence flake formation on a flake-by-flake basis and that reflect decisions made by knappers about platform selection, preparation, and core-surface management. We explore the broader potential of our model with a cross-validation approach, and we describe a means of assessing flake form on a continuum wherein variability among assemblages separated by large expanses of space and time can be meaningfully explored.

New horizons in reconstructing past human behavior: Introducing the "Tübingen University Validated Entheses-based Reconstruction of Activity" method.

An accurate reconstruction of habitual activities in past populations and extinct hominin species is a paramount goal of paleoanthropological research, as it can elucidate the evolution of human behavior and the relationship between culture and biology. Variation in muscle attachment (entheseal) morphology has been considered an indicator of habitual activity, and many attempts have been made to use it for this purpose. However, its interpretation remains equivocal due to methodological shortcomings and a paucity of supportive experimental data. Through a series of studies, we have introduced a novel and precise methodology that focuses on reconstructing muscle synergies based on three-dimensional and multivariate analyses among entheses. This approach was validated using uniquely documented anthropological samples, experimental animal studies, histological observations, and geometric morphometrics. Here, we detail, synthesize, and critically discuss the findings of these studies, which overall point to the great potential of entheses in elucidating aspects of past human behavior.

GPS-identified vulnerabilities of savannah-woodland primates to leopard predation and their implications for early hominins.

Predation is thought to have been a key selection pressure in primate evolution, especially in the savannah-woodland habitats where several early hominin species lived. However, predator-primate prey relationships are still poorly understood because human presence often deters predators, limiting our ability to quantify the impact of predation. Synchronized high-resolution tracking of leopards (Panthera pardus), vervets (Chlorocebus pygerythrus), and olive baboons (Papio anubis) during a 14-month study in Kenya revealed that increased vulnerability to leopard predation was not associated with higher encounter rates, smaller body size, smaller group size, or greater distance from refuges, contrary to long-standing inferences. Instead, the initiation, rate, timing, and duration of encounters, outcome of approaches, and predation events showed only a diel pattern of differential vulnerability. In the absence of human observers, vervets were more vulnerable during the day, whereas baboons were more vulnerable at night, but overall neither species was more vulnerable than the other. As our results show that leopards avoided baboons during the day and hunted them at night, we suggest that the same pattern would have applied to hominins-because they were even larger than baboons and bipedal, resulting in similarly offensive capability on the ground during the day but poorer agility in the trees at night, especially as they became committed bipeds. Drawing from hominid behavior and archaeopaleontological and ethnographic evidence, we hypothesize that ground-sleeping hominins initially dealt with this formidable threat by using stone tools to modify Acacia branches into 'bomas', thorny enclosures that provided nighttime shelter. The ability of hominins to create their own nightly refuges on the ground wherever Acacia spp. were available would have allowed them to range more widely, a crucial step in furthering the spread of hominins across Africa and beyond.

Consistent C3 plant habitat of hominins during 400-300 ka at the Longyadong Cave site (Luonan Basin, central China) revealed by stable carbon isotope analyses of loess deposits.

The proportions of woody and grassland taxa in terrestrial ecosystems played an important role in the origin and evolution of early Palaeolithic hominins. However the influence of ecosystem changes on hominin behavior and adaptations in Asia has not been studied in detail. Hominins have exploited the Luonan Basin in the Eastern Qinling Mountains, central China, since the early Paleolithic. Dated sites, consisting of alternating loess and soil deposits with in situ artefacts, are common in the region, and provide a detailed record of Early to Middle Pleistocene hominin environments. Here, we present the results of measurements of the stable carbon isotopic composition of soil organic matter (δ13CSOM) in the loess-paleosol sequences from the Longyadong Cave site. Our analyses of δ13CSOM show that for at least 400 ka the Longyadong Cave site and its surroundings were dominated by C3 woody plants, whereas the nearby Liuwan site was dominated by C4 and C3 mixed grassland or woody grassland vegetation. These findings demonstrate that between 400 and 300 ka in the Luonan Basin, hominins occupied a habitat consisting of a mosaic of grassland and woodland/forest. Although the vegetation of the region changed in response to the glacial-interglacial climatic cycles, patches of woody vegetation in landscapes such as at Longyadong Cave site persisted continuously. Such environments seem to be have been favored by hominins living in the Luonan Basin, possibly because they provided a diverse range of food resources during both glacial and interglacial intervals of the Middle Pleistocene, when most of northern China was experiencing an increasing trend of drying and cooling and steppe environments were expanding. Thus, the Luonan Basin would have served as a refugium for hominin occupation in China during the Middle Pleistocene.

Trabecular architecture in the thumb of Pan and Homo: implications for investigating hand use, loading, and hand preference in the fossil record.

OBJECTIVES: Humans display an 85-95% cross-cultural right-hand bias in skilled tasks, which is considered a derived behavior because such a high frequency is not reported in wild non-human primates. Handedness is generally considered to be an evolutionary byproduct of selection for manual dexterity and augmented visuo-cognitive capabilities within the context of complex stone tool manufacture/use. Testing this hypothesis requires an understanding of when appreciable levels of right dominant behavior entered the fossil record. Because bone remodels in vivo, skeletal asymmetries are thought to reflect greater mechanical loading on the dominant side, but incomplete preservation of external morphology and ambiguities about past loading environments complicate interpretations. We test if internal trabecular bone is capable of providing additional information by analyzing the thumb of Homo sapiens and Pan. MATERIALS AND METHODS: We assess trabecular structure at the distal head and proximal base of paired (left/right) first metacarpals using micro-CT scans of Homo sapiens (n = 14) and Pan (n = 9). Throughout each epiphysis we quantify average and local bone volume fraction (BV/TV), degree of anisotropy (DA), and elastic modulus (E) to address bone volume patterning and directional asymmetry. RESULTS: We find a right directional asymmetry in H. sapiens consistent with population-level handedness, but also report a left directional asymmetry in Pan that may be the result of postural and/or locomotor loading. CONCLUSION: We conclude that trabecular bone is capable of detecting right/left directional asymmetry, but suggest coupling studies of internal structure with analyses of other skeletal elements and cortical bone prior to applications in the fossil record.

Taphonomy of fossils from the hominin-bearing deposits at Dikika, Ethiopia.

Two fossil specimens from the DIK-55 locality in the Hadar Formation at Dikika, Ethiopia, are contemporaneous with the earliest documented stone tools, and they collectively bear twelve marks interpreted to be characteristic of stone tool butchery damage. An alternative interpretation of the marks has been that they were caused by trampling animals and do not provide evidence of stone tool use or large ungulate exploitation by Australopithecus-grade hominins. Thus, resolving which agents created marks on fossils in deposits from Dikika is an essential step in understanding the ecological and taphonomic contexts of the hominin-bearing deposits in this region and establishing their relevance for investigations of the earliest stone tool use. This paper presents results of microscopic scrutiny of all non-hominin fossils collected from the Hadar Formation at Dikika, including additional fossils from DIK-55, and describes in detail seven assemblages from sieved surface sediment samples. The study is the first taphonomic description of Pliocene fossil assemblages from open-air deposits in Africa that were collected without using only methods that emphasize the selective retention of taxonomically-informative specimens. The sieved assemblages show distinctive differences in faunal representation and taphonomic modifications that suggest they sample a range of depositional environments in the Pliocene Hadar Lake Basin, and have implications for how landscape-based taphonomy can be used to infer past microhabitats. The surface modification data show that no marks on any other fossils resemble in size or shape those on the two specimens from DIK-55 that were interpreted to bear stone tool inflicted damage. A large sample of marks from the sieved collections has characteristics that match modern trampling damage, but these marks are significantly smaller than those on the DIK-55 specimens and have different suites of characteristics. Most are not visible without magnification. The data show that the DIK-55 marks are outliers amongst bone surface damage in the Dikika area, and that trampling is not the most parsimonious interpretation of their origin.