Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik‐Tash child

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.     

Vindija cave and the modern human peopling of Europe

Vindija cave in Croatia has yielded the youngest securely dated Neandertal skeletal remains in Central/Eastern Europe. In addition, these remains have been found in association with archaeological material exhibiting Upper Paleolithic elements. Due to its geographic location and date, the Vindija remains are particularly crucial for the understanding of initial modern human peopling of Europe and the nature of the Neandertal demise. The significance of archaeological and paleontological finds and hominin fossils from this site is discussed in the light of new finds at Vindija and recent developments in the fields of paleoanthropology and prehistoric archaeology. Furthermore, the impact of revised chronology for several crucial specimens and sites throughout Europe, including Vindija, is discussed.     

A modern human humerus from the early aurignacian of Vogelherdhöhle (Stetten, Germany)

Implicit in much of the discussion of the cultural and population biological dynamics of modern human origins in Europe is the assumption that the Aurignacian, from its very start, was made by fully modern humans. The veracity of this assumption has been challenged in recent years by the association of Neandertal skeletal remains with a possibly Aurignacian assemblage at Vindija Cave (Croatia) and the association of Neandertals with distinctly Upper Paleolithic (but non-Aurignacian) assemblages at Arcy-sur-Cure and St. C?esaire (France). Ideally we need human fossil material that can be confidently assigned to the early Aurignacian to resolve this issue, yet in reality there is a paucity of well-provenanced human fossils from early Upper Paleolithic contexts. One specimen, a right humerus from the site of Vogelherd (Germany), has been argued, based on its size, robusticity, and muscularity, to possibly represent a Neandertal in an Aurignacian context. The morphological affinities of the Vogelherd humerus were explored by univariate and multivariate comparisons of humeral epiphyseal and diaphyseal shape and strength measures relative to humeri of Neandertals and Early Upper Paleolithic (later Aurignacian and Gravettian) modern humans. On the basis of diaphyseal cross-sectional geometry, deltoid tuberosity morphology, and distal epiphyseal morphology, the specimen falls clearly and consistently with European early modern humans and not with Neandertals. Along with the other Vogelherd human remains, the Vogelherd humerus represents an unequivocal association between the Aurignacian and modern human morphology in Europe.     

New discoveries and interpretations of hominid fossils and artifacts from Vindija Cave, Croatia

Beginning with excavations during the 1970s, Vindija Cave (Croatia) has yielded significant Middle and Upper Paleolithic fossil and archaeological finds. We report on seven recently identified hominid fossils, a newly associated partial hominid cranial vault from level G(3), nine possible bone retouchers, and a revised interpretation of the Mousterian artifact assemblage from the site. This new information reinforces our knowledge of the complex biocultural phenomena revealed in unit G and earlier deposits at Vindija. Six of the new hominid fossils derive from stratigraphic units G and I, while one lacks exact provenience. All specimens preserving diagnostic anatomy are from Neandertals. One of the postcranial remains, a radius fragment which exhibits Neandertal-like anatomy, comes from level G(1)and is congruent with the previously established association of Neandertals with an early Upper Paleolithic industry at the site. The partial cranial vault represents the most complete Neandertal from Vindija. The possible retouchers derive from unit G. Our analysis of these artifacts suggests that both percussion and pressure techniques may have been used by Neandertals in the final stage of tool production (retouching). This paper also presents a revision of the artifact analysis for late Mousterian level G(3). We separated raw materials into two main groups due to the differing ways that the materials fracture and the differing morphology of the debitage. The use of raw material in level G(3)is different from earlier Middle Paleolithic levels at Vindija. This indicates that the G(3)late Neandertals were making choices regarding source material somewhat more like the Upper Paleolithic people at the site. When interpreted within a larger regional framework, the Vindija archaeological and hominid fossil remains demonstrate a complex, mosaic pattern of biocultural change in the Late Pleistocene of south-central Europe.     

A critical review of the German Paleolithic hominin record

We review the hominin fossil record from western Central Europe in light of the recent major revisions of the geochronological context. The mandible from Mauer (Homo heidelbergensis), dated to circa 500,000 years ago, continues to represent the earliest German hominin and may coincide with the occupation of Europe north of the high alpine mountain chains. Only limited new evidence is available for the Middle Pleistocene, mostly in the form of skull fragments, a pattern that may relate to taphonomic processes. These finds and their ages suggest the gradual evolution of a suite of Neandertal features during this period. Despite new finds of classic Neandertals, there is no clear proof for Neandertal burial from Germany. Alternatively, cut marks on a skull fragment from the Neandertal type site suggest special treatment of that individual. New Accelerator Mass Spectrometry (AMS) radiocarbon dates of previous finds leave little reliably dated evidence for anatomically modern humans (AMH) in Europe before 30,000 BP; the remains from Hahn?fersand, Binshof-Speyer, Paderborn-Sande, and Vogelherd are now of Holocene age. Thus, a correlation of AMH with the Aurignacian remains to be proven, and the general idea of a long coexistence of Neandertals and AMH in Europe may be questioned. In western Central Europe, evidence of Gravettian human fossils is also very limited, although a new double grave from lower Austria may be relevant. The only dated burial from the German Upper Paleolithic (from Mittlere Klause) falls into a time period (circa 18,600 BP) represented by only a few occupation sites in western Central Europe. A number of human remains at Magdalenian sites appear to result from variable (secondary) burial practices. In contrast, the Final Paleolithic (circa 12,000-9600 cal. BC) yields an increase of hominin finds, including multiple burials (Bonn-Oberkassel, Neuwied-Irlich), similar to the situation in western and southern Europe.     

Dental remains from the Grotte du Renne at Arcy-sur-Cure (Yonne)

Human remains associated with the earliest Upper Paleolithic industries are sparse. What is preserved is often fragmentary, making it difficult to accurately assign them to a particular species. For some time it has been generally accepted that Neandertals were responsible for the Chatelperronian and anatomically modern humans for the early Aurignacian industries. However, the recent re-dating of several of the more-complete modern human fossils associated with the early Aurignacian (e.g., Vogelherd) has led some to question the identity of the makers and the context of these early Upper Paleolithic industries. The Grotte du Renne at Arcy-sur-Cure, France has yielded many hominin remains, from Mousterian, Chatelperronian, Aurignacian, and Gravettian layers. Previously, a child's temporal bone from the Chatelperronian Layer Xb was recognized as belonging to a Neandertal; however, most of the teeth from Chatelperronian layers VIII-X remain unpublished. We describe the dental remains from the Chatelperronian layers, place them in a comparative (Mousterian Neandertal and Upper Paleolithic modern human) context, and evaluate their taxonomic status. The teeth (n = 29) represent a minimum of six individuals aged from birth to adult. The permanent dental sample (n = 15) from the Chatelperronian layers of Arcy-sur-Cure exhibits traits (e.g., lower molar mid-trigonid crest) that occur more frequently in Neandertals than in Upper Paleolithic modern humans. Furthermore, several teeth show trait combinations, including Cusp 6/mid-trigonid crest/anterior fovea in the lower second molar, that are rare or absent in Upper Paleolithic modern humans. The deciduous teeth (n = 14) significantly increase the sample of known deciduous hominin teeth and are more similar to Mousterian Neandertals from Europe and Asia than to Upper Paleolithic modern humans. Thus, the preponderance of dental evidence from the Grotte du Renne strongly supports that Neandertals were responsible for the Chatelperronian industry at Arcy-sur-Cure.     

The current state of korean paleoanthropology

The hominid fossil and Paleolithic archaeology records from the Korean Peninsula are extensive, but relatively little is known about the Korean human evolutionary record outside this region. The Korean paleoanthropological record is reviewed here in light of major research issues, including the hominid fossil record, relative and chronometric dating, lithic analysis, hominid subsistence, and the presence of bone tools, art and symbolism. Some of the major conclusions drawn from this review include: (1) hominid fossils have been found in nine separate sites on the Korean Peninsula; (2) possible Homo erectus fossils are present in North Korea; (3) Ryonggok Cave, in North Korea, has exposed the remains of at least five archaic Homo sapiens individuals; (4) a possible burial of an anatomically modern Homo sapiens child, discovered in Hungsu Cave in South Korea, has been tentatively dated to roughly 40,000 years ago; (5) handaxes and cleavers have been found at a number of sites near Chongokni and they appear to date to at least 100,000 years ago; and (6) taphonomic studies are necessary for addressing issues related to determining the nature of hominid-carnivore interaction over similar resources (e.g. carcasses and shelter); and the presence/absence of Early Paleolithic bone tools, art, and symbolism in Korea.     

Re-analysis of the hominid radii from Cave of Hearths and Klasies River Mouth, South Africa

Two of the few postcranial fragments from the late Early Stone Age and/or the Middle Stone Age of southern Africa are the proximal radii from the Cave of Hearths and Klasies River Mouth. The Cave of Hearths fossil is metrically indistiguishable from both archaic (e.g., Neandertals) and recent humans, and presents a mosaic of primitive and modern features. The primitive include a relatively slender neck and thick cortical bone (the latter of which distinguishes recent humans from archaic, Early Modern, and Upper Paleolithic hominids); the modern includes an anteromedially (rather than medially) facing radial tuberosity. Its extreme collo–diaphyseal angle is unusual, although it can be matched by modern homologues. The neck–shaft angle of some Neandertal and Early Modern radii also appears to match that of the Cave of Hearths specimen. The Klasies River Mouth radius also has thick cortical bone of the neck. It is morphologically indistinguishable from Early Modern and Neandertal homologues. These, and other fossils, suggest a mosaic pattern of evolution in the postcranial skeleton of the late Early Stone Age and/or Middle Stone Age inhabitants of sub-Saharan Africa.     

The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW,NW Germany)

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100 m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave.     

Into Eurasia: A geometric morphometric re-assessment of the Upper Cave (Zhoukoudian) specimens

Since the discovery of the human remains from the Upper Cave of Zhoukoudian in the 1930s there has been speculation over their affinities. In particular, the degree to which the three adult crania exhibit recent East Asian morphology, as well as their degree of within-group variability, has long been debated. Several early researchers described a resemblance to East Asian populations, but these findings have been for the most part rejected by more recent metric and non-metric analyses. Nevertheless, the Upper Cave specimens have not been classified conclusively into any recent modern human population to which they have been compared, and classification results differ for each cranium. Here, the question of the affinities of Upper Cave 101 and 103, the two better-preserved crania, is examined from the perspective of the Late Pleistocene human fossil record using the methodology of 3-D geometric morphometrics. The degree of morphological variation between the two specimens is also evaluated within the context of recent population variability. Neurocranial and facial morphology are analyzed separately so as to maximize comparative samples. Results show a morphological resemblance of the Upper Cave material to Upper Paleolithic Europeans. It is proposed that the Upper Cave specimens retain important aspects of modern human ancestral morphology, and possibly share a recent common ancestral population with Upper Paleolithic Europeans, in accordance with the Single Origin model of modern human origins.     

Hovk 1 and the Middle and Upper Paleolithic of Armenia: a preliminary framework

The territory of present day Armenia is a geographic contact zone between the Near East and the northern Caucasus. Armenian Middle and Upper Paleolithic records are both few and patchy as a result of the historical paucity of systematic archaeological research in the country. Consequently, it is currently difficult to correlate the Armenian Middle and Upper Paleolithic records with those from other neighboring regions. We present new archaeological and chronometric data (luminescence, U-Th, and ¹⁴C) from our ongoing research at Hovk 1 Cave in northeast Armenia. We discuss in particular two activity phases in Hovk 1 Cave for which we have outline chronometric data: (1) an early Middle Paleolithic occupational phase, dated by optically stimulated luminescence (OSL) to 104 ± 9.8 ka BP_OSL; and (2) a Paleolithic occupational phase characterized by microlithic flakes dated by AMS ¹⁴C to 39,109 ± 1,324 calibrated years BP_Hulu. The two phases are separated by a hiatus in hominin occupation corresponding to MIS 4 and an episode in early MIS 3. These chronometric data, taken together with the preliminary paleoenvironmental reconstruction of the Hovk 1 Cave and environment, suggest that these activity phases represent short-lived and seasonal use of the cave presumably by small groups of hunters during episodes of mild climate. Neither tool manufacture nor butchery appears to have taken place within the cave, and consequently, the archaeological record included, for the most part, finished tools and blanks. We address the chronology and techno-typological aspects of Hovk 1 lithics in relation to: (1) the Paleolithic records of Armenia, and (2) the broader interregional context of early Middle Paleolithic hominin occupation and the Middle-Upper Paleolithic transition in the Caucasus.     

Os fossiles humains des grottes Muierii et Cioclovina, Roumanie

This paper presents the cultural and archaeological context of the human fossil bones from Muierii Cave, dated by us to the age of 30 150 ± 800 ¹⁴C years BP (Before Present) or 34 810 ± 927 cal years BP (calibrated years Before Present), and from Cioclovina Cave, dated to the age of 29 000 ± 700 ¹⁴C years BP or 33 540 ± 832 cal years BP, in the Southern Carpathians. These are among the most ancient dated human fossil remains from Central and South-Eastern Europe and are described in conjunction with other sites with Mousterian assemblages of the recent Neanderthal population, and sites with Aurignacian assemblage of early modern humans, from Romanian region, for the interval of time 34,000-26,000, the transitional period from the Middle Paleolithic to the Upper Paleolithic.     

Fossil human remains from Bolomor Cave (Valencia, Spain)

Systematic excavations carried out since 1989 at Bolomor Cave have led to the recovery of four Pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. All of these specimens date to the late Middle and early Late Pleistocene (MIS 7-5e). The fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-Homo sapiens) members of the genus Homo. Among the dental remains, the lack of a midtrigonid crest in the M₁ represents a departure from the morphology reported for the majority of Neandertal specimens, while the large dimensions and pronounced shoveling of the marginal ridges in the C¹ are similar to other European Middle and late Pleistocene fossils. The parietal bone is very thick, with dimensions that generally fall above Neandertal fossils and resemble more closely the Middle Pleistocene Atapuerca (SH) adult specimens. Based on the presence of archaic features, all the fossils from Bolomor are attributed to the Neandertal evolutionary lineage.     

Neutron microtomography‐based virtual extraction and analysis of a cercopithecoid partial cranium (STS 1039) embedded in a breccia fragment from sterkfontein member 4 (South Africa)

The Plio‐Pleistocene karstic sedimentary deposits of Sterkfontein Cave, South Africa, yielded numerous fossil primate specimens embedded in blocks of indurated breccia, including the partial cercopithecoid cranium labelled STS 1039. Because the surrounding matrix masks most of its morphology, the specimen remains taxonomically undetermined. While the use of X‐ray microtomography did not allow extracting any structural information about the specimen, we experimented a new investigative technique based on neutron microtomography. Using this innovative approach, we successfully virtually extracted, reconstructed in 3D and quantitatively assessed the preserved dentognathic structural morphology of STS 1039, including details of its postcanine maxillary dentition. Following comparative analyses with a number of Plio‐Pleistocene and extant cercopithecoid taxa, we tentatively propose a taxonomic attribution to the taxon Cercopithecoides williamsi. Our experience highlights the remarkable potential of this novel imaging method to extract diagnostic information and to identify the fossil remains embedded in hard breccia from the South African hominin‐bearing cave sites. Am J Phys Anthropol 159:737–745, 2016. © 2015 Wiley Periodicals, Inc.     

Identifying the morphological signatures of hybridization in primate and human evolution

Recent studies point to contact and possible admixture among contemporaneous hominin species during the Plio-Pleistocene. However, detection of hybridization in fossils-and especially fossil hominins-is contentious, and it is hindered in large part by our lack of understanding about how morphological hybridity is manifested in the primate skeleton. Here, we report on a study of known-pedigree, purebred yellow and olive baboons (n = 112) and their hybrids (n = 57), derived from the baboon colony of the Southwest Foundation for Biomedical Research. The hybrids were analyzed in two different groups: (1) F1 = olive x yellow first-generation hybrids; (2) B1 = olive x F1 backcross hybrids. Thirty-nine metric variables were tested for heterosis and dysgenesis. Nonmetric data were also collected from the crania. Results show that these primate hybrids are somewhat heterotic relative to their parental populations, are highly variable, and display novel phenotypes. These effects are most evident in the dentition and probably indicate the mixing of two separately coadapted genomes and the breakdown in the coordination of early development, despite the fact that these populations diverged fairly recently. Similar variation is also observed in museum samples drawn from natural hybrid zones. The results offer a strategy for detecting hybrid zones in the fossil record; implications for interpreting the hominin fossil record are discussed.     

Out of the North Sea: the Zeeland Ridges Neandertal

In 2001, a portion of human frontal bone was discovered in sediments extracted from the bottom of the North Sea, 15 km off the coast of the Netherlands. The extraction zone is located in the so-called Zeeland Ridges area located at 51°40′ northern latitude and 3°20′ eastern longitude. The specimen was dredged up from sediments containing Late Pleistocene faunal remains and Middle Palaeolithic artefacts, including well-finished small handaxes and Levallois flakes. The details of the supraorbital morphology, as well as the quantitative assessment of the shape of the external surface of the squama using traditional and 3D geometric morphometrics, unambiguously assign the Zeeland Ridges frontal bone to Homo neanderthalensis. Carbon and nitrogen isotopic analysis indicate that the Zeeland Ridges hominin, like other Neandertals, was highly carnivorous and does not show evidence for the consumption of aquatic foods. A lesion on the outer table and diploic layer of the bone in the area of the supratoral sulcus can be interpreted as the result of an intradiploic epidermoid cyst, a type of neoplasm diagnosed for the first time in Neandertal remains. So far, the Zeeland Ridges Neandertal is the first Pleistocene fossil hominin found under seawater and the first recorded in the Netherlands.     

A new hominin from the Basal Member of the Hadar Formation, Dikika, Ethiopia, and its geological context

In this paper we report for the first time hominin remains from the Basal Member of the Hadar Formation at Dikika, in the Awash Valley of Ethiopia, dating to greater than 3.4 Ma. The new fossil, DIK-2-1, is a fragment of a left mandible and associated dentition. The mandible is attributed to Australopithecus afarensis. However, the new fossil exhibits some metric and morphological features that have not previously been seen in the A. afarensis hypodigm, increasing the already impressive degree of variation in the mandibular sample of the species.