Human Evolution, Homo heidelbergensis

The Middle Pleistocene (~721-126 thousand-years-ago) is one of the most complex and unclear periods with regards to the evolutionary history of genus Homo. Remains from Africa, Europe, and sometimes Asia are assigned to “Homo heidelbergensis”: a species theoretically defined around the ~600,000-year-old Mauer mandible. However, many specimens do not align closely with the holotype, and the apparently-broad polymorphism during this period, combined with the emergence of modern human, Neanderthal, and Denisovan lineages have made the designation H. heidelbergensis extremely controversial.

A geometric-morphometric analysis of posterior dental crowns was undertaken, with the aim of quantifying and assessing levels of intra-specific shape variability within H. heidelbergensis in comparison with other taxa. The results showed that Middle Pleistocene specimens generally exhibited larger inter-individual shape variation than other hominin species and may include several palaeodemes, providing further support to the illegitimacy of H. heidelbergensis. However, it may not necessarily be concluded from this study that the specimens represent more than one broad, over-arching species; at least, within this sample of mostly-European specimens. Alternatively, it may provide support for broad intra-taxic variability, and leaves the status of H. heidelbergensis still to be clarified. Though many interpretations of H. heidelbergensis are found in palaeoanthropological literature, it may not necessarily be inaccurate as a cross-continental species lasting from ~700kya until speciation events, ~250kya.

The addition of Afro-Eurasian specimens would provide a fairer assessment of H. heidelbergensis as a cross-continental taxon. Furthermore, this study adds to the lack of clarity with regards to acceptable levels of polymorphism within the genus Homo.

The debate over whether Africa was the sole home of hominin species (excepting Homo neanderthalensis and Homo floresiensis) is not completely won (Dennell, 2009: 466). Homo erectus (sensu lato) is often thought to be the first species to have left Africa and entered Asia (Langbroek, 2004: 11; Klein, 2005: 102), with occasional discussions on earlier hominids (such as australopithecines) having made this symbolic breakthrough (Templeton, 2002: 48; Dennell and Roebroeks, 2005: 1100). The role of climate has often been used to discuss the emergence of a particular species, however it has been less discussed when approaching why a species – (Homo erectus) – may have evolved into a new species – (Homo heidelbergensis) – on one continent, whilst seemingly persisting for a large period of time on another, with little change to its skeletal morphology.
Homo erectus became ubiquitous (Bonnefille, 2010: 408) across a number of environments across Africa and Asia, from high plateau to sea level, temperate to tropical, desert to rainforest as well as persisting from c.1.7 Ma to maybe c. 60 Ka (Klein, 2005: 106). The emergence of this species is believed to have been triggered by climatic changes c. 1.8 Ma, as was its dispersal across the old world (or before) (Bar Yosef, 1998: 267; Clark, 1998: 437). Ultimately, the equation for hominin occupation and survival across the earth required access to water, food and stone (Dennell, 2009: 476).

Western Eurasia yielded a rich Middle (MP) and Late Pleistocene (LP) fossil record documenting the evolution of the Neandertals that can be analyzed in light of recently acquired paleogenetical data, an abundance of archeological evidence, and a well-known environmental context. Their origin likely relates to an episode of recolonization of Western Eurasia by hominins of African origin carrying the Acheulean technology into Europe around 600 ka. An enhancement of both glacial and interglacial phases may have played a crucial role in this event, as well as in the subsequent evolutionary history of the Western Eurasian populations. In addition to climatic adaptations and an increase in encephalization, genetic drift seems to have played a major role in their evolution. To date, a clear speciation event is not documented, and the most likely scenario for the fixation of Neandertal characteristics seems to be an accretion of features along the second half of the MP. Although a separation time for the African and Eurasian populations is difficult to determine, it certainly predates OIS 11 as phenotypic Neandertal features are documented as far back as and possibly before this time. It is proposed to use the term ''Homo rhodesiensis'' to designate the large-brained hominins ancestral to H. sapiens in Africa and at the root of the Neandertals in Europe, and to use the term ''Homo neanderthalensis'' to designate all of the specimens carrying derived metrical or non-metrical features used in the definition of the LP Neandertals.

"This thesis aims to explore whether idiosyncrasies in Acheulean handaxe manufacture can be seen and, if so, whether these can be used to trace the actions of hominins within the Lower Palaeolithic. This analysis has important implications for the application of current social theory to Palaeolithic contexts, which advocates a 'bottom-up' approach to archaeological study. This socially orientated theoretical approach emphasises the individual as the primary unit of analysis. However, as Hopkinson and White (2005) state, there is currently no methodology for such an analysis, rendering many discussions as exercises in what has been termed 'theoretical storytelling'.

Using a series of innovative experiments, the questions of whether the individual is a viable unit of analysis was tested. The results show that a suite of other factors that also contribute to stone tool manufacture currently masks the actions of individuals. Chief amongst these is the variability in the raw material nodules selected for reduction. However, intra-site variability may indicate differences that are linked to socially mediated knapping strategies, or 'group templates' (c.f. Ashton and McNabb 1994). While this possibility requires further exploration, this thesis suggests that the individual is currently not viable as a primary unit of analysis within Palaeolithic archaeology and stresses that the theories posited from the standpoint of the individual cannot be interpreted as fact. At the same time, it appears that further work needs to be conducted that focuses on the more traditional group as the primary analytical unit and the prospect of teasing apart the interplay between individuals, groups and the effects of raw material variability."

Volume II of this thesis can be accessed via http://etheses.dur.ac.uk/5946/

Definition: A controversial Middle Pleistocene (~780-130 ka) hominin species, part of the same genus (Homo) as extant humans. Introduction: The taxonomy and phylogeny of Homo heidelbergensis is much debated (for a review see Stringer, 2012), so much so that this period in human evolution has been dubbed the 'muddle in the Middle [Pleistocene]'. There is little agreement on which specimens should be included in H. heidelbergensis (that is to say, the group of specimens which constitute what is known as its 'hypodigm'), nor even whether it constitutes a valid species at all. Many Middle Pleistocene hominins (taxa more closely related to H. sapiens than to chimpanzees) share primitive features with earlier species, such as H. erectus, but also share derived traits with later Pleistocene specimens such as H. sapiens and Neanderthals (e.g., Rightmire, 2013). This mosaic morphology has led researchers to group them together as a single Afro-European species designation: H. heidelbergensis. Some have also suggested that additional Asian Middle Pleistocene fossils may extend the range of H. heidelbergensis (e.g., Stringer, 2012). One reason for the great interest in H. heidelbergensis is that the Middle Pleistocene is the epoch during which the H. sapiens lineage split from our sister taxa, the Neanderthals and Denisovans and H. heidelbergensis has long been considered a possible last common ancestor (LCA) for these two lineages (Buck & Stringer, 2014; Mounier et al., 2009; Rightmire, 2008; Stringer, 2012). Knowing the identity of the LCA would allow us to see which differences between our own species and Neanderthals are derived in which lineage, enabling us to better understand our own evolution and that of our closest relatives.

Hominin taxonomies distinguish taxa by criteria of time, space, and morphology. These »species« are defined typologically and described using a mixture of primitive and derived characters. The adoption of a cladistic perspective has led to an extension of phylogenetic principles to the problem of taxon definition, and often resulted in the simple recasting of these descriptive characters as apomorphies of their respective taxa. However, characters cannot be identified as apomorphies in the absence of a well- supported phylogenetic hypothesis for these taxa.
The present study is a phylogenetic analysis of Middle to Late Pleistocene fossil specimens assigned to Homo antecessor, H. heidelbergensis, H. neanderthalensis, and H. sapiens, employing characters that are con- sidered taxonomically significant for these groups. Craniodental specimens were examined and scored for a maximum of 77 discrete and continuous characters. Phylogenetic analyses employing a variety of character and taxon matrices, coding schemes, and outgroup taxa were conducted using PAUP* and MacClade, with the resulting phylogenies and synapomorphies supporting clades compared to current systematic hypotheses concerning these fossils.
Neandertals and modern H. sapiens both formed consistent, well-sup- ported clades. H. heidelbergensis was not monophyletic in the results of any analyses and is most likely a grade taxon. There was no support for the identification of H. antecessor as the unique ancestor of modern humans. Very few traditional »taxonomic characters« support the clades they are meant to describe. These results suggest that paleoanthropologists should be more cautious about their identification of species in the fossil record and phylogenetic hypotheses concerning these taxa.

La quantité importante de niveaux d'occupations humaines identifiées dans le remplissage de la Caune de l'Arago, daté de 700 000 ans à la base et de 100 000 ans au sommet, et leur exceptionnelle richesse, ont donné lieu à une étude des types d'habitat et des modes de vie des chasseurs paléolithiques entre 600 000 ans et 400 000 ans. Les résultats des études pluridisciplinaires menées depuis de nombreuses années sur ces niveaux archéologiques permettent aujourd'hui de proposer une typologie de l'habitat en grotte pour le Pléistocène moyen du sud de la France. En effet, quatre types d'habitat ont pu être identifiés et décrits. Ils ont été définis à partir de différents caractères comme l'épaisseur de l'accumulation archéologique, les espèces animales représentées, les saisons d'abattages identifiées, l'état de surface des ossements et leur conservation, les caractéristiques morphotechniques et typologiques des assemblages lithiques, les catégories de matières premières minérales utilisées et le territoire d'exploitation défini à partir de celles-ci. Ces quatre types d'habitat sont : l'habitat de longue durée, où la grotte a été utilisée comme camp résidentiel de base, le campement temporaire saisonnier, où la grotte est utilisée comme campement secondaire, la halte de chasse et le bivouac où la grotte est cette fois considérée comme un refuge pour un séjour de courte durée. Les modes de vie sont étroitement liés au type d'habitat et sont généralement tous orientés vers une activité principale : la chasse. Les activités sont diversifiées dans les occupations de longues durées et saisonnières alors qu'elles sont plus réduites et très spécialisées dans les occupations brèves. En résumé la Caune de l'Arago présente toutes les caractéristiques d'un site à usage multiple qui a été utilisé à de nombreuses reprises par des groupes de chasseurs ou des groupes familiaux d'anténéandertaliens qui ont laissé de nombreuses traces de leurs activités nous permettant de connaître leurs habitats et leurs modes de vie.Several human occupations have been identified within the deposits of the Caune de l’Arago Cave site, dated from 700 000 years to 100 000 years old. The exceptional wealth of these archeological levels provides an opportunity for the study of different types of habitat and behavioral patterns of Paleolithic hunters between 600 000 and 400 000 years ago. Results from preliminary studies, carried out over the last 30 years on these archeological levels, allow for the proposition of a typology for cave habitats during the Middle Pleistocene in the south of France. In fact, four types of habitat were identified and defined. They have been described according to different characteristics such as, the thickness of the archeological accumulation, the animal species represented, the season during which the animals were hunted, the state of conservation of the fossils discovered, the morpho-technological and typological characteristics of the lithics, the categories of raw materials used and the territory exploited by the hominids inhabiting the cave, as defined by the raw materials represented. The four habitation types defined are: the long duration habitat, where the cave was used as a home base; the temporary seasonal habitat, in which case the cave served as a secondary campsite; the hunting stopover and the bivouac, during which the cave was used as a refuge for short term stays. Behavioral patterns appear to be directly related to the type of habitat and oriented towards a principal activity: hunting. The activities performed by the inhabitants of the cave seem to be more diversified during prolonged or seasonal occupations, whereas they appear to be reduced or very specialised during brief stays. Generally, The characteristics studied show that the Caune de l’Arago Cave site had multiple uses over time for Anteneandertalian family groups and hunters who left traces of their activities, allowing us to better understand their lifestyle.

In 1995-1996 two isolated hominin lower incisors were found at the middle Pleistocene site of Boxgrove
in England, with Lower Palaeolithic archaeology. Boxgrove 2 is a permanent lower right central incisor
and Boxgrove 3 a permanent lower left lateral incisor. They were found separately, but close to one
another and appear to belong to the same individual. The Boxgrove 1 tibia discovered in 1993 came from
a different stratigraphic context and is thus believed to represent a different individual. This paper
describes the morphology of the incisors, which is similar to other middle Pleistocene hominin specimens
and, as with the tibia, suggests that they could be assigned to Homo heidelbergensis (recognising
that the taxonomic status of this species is still a matter of debate). The incisors show substantial
attrition associated with secondary dentine deposition in the pulp chamber and clearly represent an
adult. They also show extensive patterns of non-masticatory scratches on the labial surfaces of both
crown and root, including some marks which may have been made postmortem. The roots were exposed
in life on their labial sides by a large dehiscence, extending almost to the root apex. This is demonstrated
by deposits of calculus, polishing, and scratching on the exposed surfaces. The dehiscence may have been
caused by repeated trauma to the gingivae or remodelling of the tooth-supporting tissues in response to
large forces applied to the front of the dentition.

The Stone Age, the longest period in the history of mankind except for the latest 5,000 years, covers a huge period of 2.6-2.7 million years, the lion share of which consists of its most ancient period - Paleolithic.
All periods of Stone Age - from the ancient Paleolithic (Oldowan) to Neolithic - are represented within the boundaries of modern Azerbaijan. Bearing Stone Age culture elements, which are typical for each period, on the monuments of Azerbaijan, they often have local distinctions illustrating on a long existence of a given culture in a given territory, with its further evolution and sometimes “export” to adjacent territories.
Complex studies of the Azykh Cave sediments make it possible to determine the chronology of the site’s habitation with a larger share of confidence. Azykh sediments strictly indicate on three differently-timed cultures of Paleolithic corresponding to the three episodes of the Cave’s being settled.
The first episode of the Cave’s settlement. The layers VII-X represent the earliest episode, no less than 1.2 million years ago in the Lower Pleistocene, perhaps, in the second half of Apsheron by the creator of the Kuruchay culture, a kind of Homo erectus; probably, the kind that was discovered in Dmanisi as well.
The second episode of the Cave’s settlement. It occurred in the Acheulian epoch. Carriers of Early Acheulian culture - layer VI - lived here in the pre-Oka and Oka period (no less than 600,000-400,000 years ago), while that of Middle Acheulian culture - in the Lihvin Glacial Age (400,000-250,000 years ago). A relatively late form of hominids resembling Homo erectus inhabited Azykh in the Middle Acheulian epoch.
When the upper sediments of layer V were being accumulated, it became colder so the inhabitants left the site (as proven by the sterile layer IV). Leaving the Cave in the period of general worsening of climate, azykhantrop unlikely “resettled” to another cave; more likely, he migrated to areas with a milder climate and for a long time because he returned to the Cave as a Neanderthal man with the early Mousterian culture after dozens of millenniums.
The third episode of the Cave’s settlement. The layer III - the latest one - was formed in the Middle Paleolithic, more exactly in Early Mousterian epoch most probably by Neanderthal people, with whom the Middle Paleolithic culture is inseparably connected.
Thus, it is possible to state 3 episodes (avoiding to indicate their duration) of the Cave’s settlement with each episode corresponding to three different Paleolithic cultures and, probably, to different kinds of Homo.

It is often difficult to differentiate between archaeological bonebeds formed by one event such as a mass kill of a single herd, and those formed by multiple events that occurred over a longer period of time. The application of high temporal resolution studies such as intra-tooth isotopic profiles on archaeological mammal cohorts offers new possibilities for exploring this issue, allowing investigators to decipher between single and multiple accumulation events.We examined 18O and 13C isotopic variations from the enamel carbonate of 23 horse third molars from the Middle Pleistocene archaeological site of Schöningen. We employed a new approach to investigate processes of fossil accumulation that uses both bulk and intra-tooth isotopic variations and takes into account animal behavior, age at death and dental development to test the degree of isotopic affinity of animals from the same fossil assemblage.
Oxygen and carbon isotope bulk values indicate that the horses from Schöningen 13 II-4 experienced relatively similar climatic and dietary regimes. Inter-individual differences of the bulk values of the horses sampled in the current study present nevertheless inter-individual variability similar to individuals from
multi-layered localities. In addition, the intra-tooth isotopic variation of specimens of the same age at death seems to indicate that the studied cohort corresponds to a mix of individuals that recorded both similar and different isotopic histories. Finally, the conditions recorded in the isotopic signal shortly before death (i.e., for teeth not fully mineralized) varied between sampled individuals, suggesting possible differences in the seasonality of death. Considering those results, we discuss the possibility that the horses from Schöningen 13 II-4 correspond to an accumulation of different death events.

The existence of calcanei in the fossil record prior to modern humans and Neandertals is very scarce. This skeletal element is fundamental to understanding the evolution of the morphology of the foot in human evolution. Here we present and metrically and comparatively describe 29 calcaneus remains from the Middle Pleistocene site of Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These calcanei belong to 15 individuals (nine adults, two adolescents and four immature individuals). The metric and morphological differences in the calcanei among Middle and Late Pleistocene hominins tend to be subtle. However, the calcanei from SH are broad and robust with large articular surfaces and most significantly, exhibit a very projected sustentaculum tali. A biomechanical and phylogenetic interpretation is proffered to explain the observed morphology of these calcanei. It has been possible to propose tentative sex assignments for the SH calcanei based on size, using methods similar to those used to establish sex from the talus bones from SH. The estimation of stature based on the calcaneus provides a mean of 175.3 cm for males and 160.6 for females, which is similar to that obtained using other skeletal parts from the site. In sum, the SH calcanei are robust with a proportionally long tubercle and a projected sustentaculum tali, which are traits shared by Neandertals.

The Northwest of Iberia has yielded one of the most complete European Middle Paleolithic records. Despite this wealth of information, very little is known about population dynamics during this period. For that reason, the main concern of this paper is to provide socio-environmental models that may help explain Early Middle Paleolithic (EMP) population dynamics in NW Iberia, assessing to what extent they were shaped by climate forces. The archaeological record is analyzed on the basis of the heuristics of ecological models, already employed in the European Pleistocene record but never at a regional scale, in order to detect long-term changes in the composition of EMP populations, and the environmental, biological and sociocultural process influencing those changes. According to the models proposed, we have detected a long-term population dynamic between MIS 11 and MIS 6, characterized by low environmental stress, high biological productivity, interaction among populations and sociocultural complexity. Eventually, this population dynamic was broken due to an extreme climate phase in late MIS 6 that had a profound impact on populations and sociocultural structures. As a result, the Upper Pleistocene population of NW Iberia was concentrated in the Cantabrian region. This area became an isolated Neanderthal glacial refugium that hosted a population with different origins and fragile long-term demographic stability.

Two skulls that are relics that would be nothing without a known provenance and an established identity. They have sparked a lot of research, some with the help of Yves Coppens who has never ceased to examine human bones in multiple forms in order to better describe their functioning and behavior since our origins, including fossils that have been found in East Africa. Anthropology needs material witnesses, and in the case which concerns us it is a question of skulls with a historical character which goes as far as a mystical plasmic presence which only exists because we know their verified history thanks to research. in laboratory. The expression is from Philip K. Dick (2012), and means that the characteristic or characters of an object or a person must be proven, identified in a historical and scientific way which in the case of Tautavel were carried out by the team of Henry de Lumley and for Mozart in forensic medicine who also discovered the cause of death

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima
de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and
three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this
study has identified unique morphological characteristics of the SH tali. They are vertically shorter than
those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet
than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the
hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a
broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene
H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for
the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for
males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.

La Sima de los Huesos, en la Sierra de Atapuerca, con más de 5.000 restos de fósiles humanos procedentes de la primera acumulación de cadáveres, supone uno de los yacimientos más importantes a nivel mundial. La gran cantidad de información que podemos extraer del estudio de ellos nos permite acercarnos aún más a nuestros antepasados y saber cómo eran, cómo vivían y cómo murieron.