The morphological evidence
Palaeoanthropologists generally have little problem seperating Neandertals and modern humans based on their gross morphologies. However, some of the earliest modern humans from central Europe have traits that have been seen as evidence for continuity between them and Neandertals. These fossils, particularly those from Peştera cu Oase in Romania and Mladeč in the Czech Republic, have been touted as exemplars for modern-Neandertal admixture. These specimens show traits that are seen in high frequencies in Neandertals, such as bunning of the occipital and the presence of a suprainiac fossa.
However, many researchers have questioned whether these traits are in fact distinctly Neandertal. For instance, the form of the occipital seems to be different in early Upper Palaeolithic populations, leading many to favour the term hemibun to describe the shape of the occipital in early Europeans. Lieberman and colleagues has gone as far as to suggest that the buns seen in these two groups are not homologous. Similarly, it has been argued that the shape of the suprainiac fossa is distinct in early modern Europeans compared to Neandertals.
A palpable difficulty in assessing proposed Neandertal traits in early modern humans is that both groups shared similar niches and some traits may be the result of lifetime behavioural adaptations or convergent evolution. Indeed, the shared robustness of these early humans is likely due to the higher physical activities of these Late Pleistocene groups than during later period.
The genetic evidence
Mitochondrial DNA (mtDNA) has some characteristics that make it ideal for analyses of ancient specimens. MtDNA is found in abundance – cells can have thousands of copies of mtDNA, while only containing two copies of nuclear DNA. Moreover, its structure and location within the cell make it more resistant to decay. All the studies of Neandertal mtDNA to date cluster outside the range for modern human mtDNA variation. However, the mitochondria contain only a small part of the total DNA that make up a genome. The possibility that Neandertal genes could show up somewhere else in the genome cannot be ruled out.
The recent announcement by Svante Pääbo that he is sure that Neandertals and modern humans had sex is quite a bold pronouncement coming from a scientist. It raises the question of whether this ascertain is based on some hard evidence they found while sequencing the Neandertal genome. It is possible that if there was some Neandertal genes passed on to the first moderns in Europe, they could have got eliminated from the subsequent gene pool as population sizes fluctuated during the more severe climatic episodes. A more likely scenario is that Pääbo’s team found evidence of modern introgression in the Neandertal genome. In all likelihood the incoming modern humans were more numerous than the Neandertals, thereby absorbing the endemic populations through genetic swamping.
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At the conference, much attention was focused on the Middle Pleistocene “muddle in the middle” , particularly the role of Homo heidelbergensis in hominin evolution. While H. heidelbergensis possesses both archaic and derived traits intermediate between H. erectus and later members of the Homo genus, it lacks uniquely derived traits or autapomorphies, which are a prerequisite for defining a species.
H. heidelbergensis has traits that have been interpreted as nascent Neandertal autapomorphies, leading some researchers to propose that there was a continuous evolution of Neandertals [4-6]. This accretion model would make H. heidelbergensis a chronospecies on the continuum of the Neandertal lineage, a view championed by Jean-Jacques Hublin. The accretion model proposes that Neandertals evolved by anagenesis, i.e. non-branching evolutionary change.
Another scenario views both the European and African H. heidelbergensis as a single species, and the last common ancestor of both Neandertals and modern humans. Alternatively, H. heidelbergensis could have become isolated in Europe and evolved into Neandertals, while the African populations led to modern humans.
During the conference, Ian Tattersall noted that while the accretion model explains some of the variation in the Middle Pleistocene, it cannot account for some outliers, such as the 28 or so specimens that have been recovered from the Sima de los Huesos in Atapuerca, Spain. Tattersall is not the first author to call the accretion model into question . Recent dates have placed the Sima fossils at just over half-a-million years old. Based on the dissimilarity between these fossils and the penicontemporaneous H. heidelbergensis from the rest of Europe, Tattersall proposes that two hominin lineages coexisted in Europe before the arrival of H. sapiens. He suggests that one line (which may include the Sima specimens) led to the Neandertals, while the branch which included H. heidelbergensis went extinct. If Tattersall is correct it would mean that the Sima fossils, which are currently classified as H. heidelbergensis, must be designated another name.
Hublin is to his guns and doesn’t see any need to reclassify the Sima material. He goes as far as to suggest binning the species name H. heidelbergensis altogether and instead reassigning all these Middle Pleistocene fossils as H. neanderthalensis. Whatever the outcome is in this debate, it appears that hominin evolution in the Middle Pleistocene is more complex than we have previously suspected.
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The gluteus maximus, which is a relatively minor muscle in quadrupeds has been reconfigured into the largest muscle in humans, in order to stabilize the pelvis and trunk in an upright position. The derived nature of the ilium of Ar. ramidus suggests that the enlargement of the gluteal maximus had already begun. The craniocaudal height of the pelvis is also reduced, which would have lowered the relatively long trunk’s centre of mass. This would have allowed for more stable bipedal locomotion.
However, the ischium is quite primitive compared to the ilia, likely to accommodate the large hindlimb musculature required for tree climbing. The two best preserved australopithicine pelves, AL 288-1 and Sts 14, both have short ischia, like those seen in modern humans. The preserved portion of the ischial ramus in Ar. ramidus is significantly larger than that found in any of the Australopithecines. A long ischium creates a greater moment arm suggesting that Ar. ramidus had relatively powerful hamstrings, a trait that is common in tree-dwelling primates.
The configuration of the ARA-VP-6/500 pelvis suggests that lower lumbars were probably posteriorly positioned, allowing for lordosis of the spine. A reduction in iliac height would have further facilitated lordosis. Lordosis positions the spine to a more forward position, so that it directly overlies the hips during erect posture. Lower spinal lordosis would have allowed the full extension of the hips and knee during extended bipedal locomotion.
Ar. ramidus was quite capable of bipedal locomotion, as attested to by the morphology of its pelvis and foot. However, its large thigh muscles and its prehensile big toe show that it was still very much adapted to arboreal life. Ar. ramidus shares arboreal adaptations that were probably present in the human-chimp last common ancestor, as well as bipedal adaptations that are so characteristic of hominins. Ar. ramidus appears to have been an arboreal ape with bipedal adaptations, rather than a biped with arboreal adaptations. It is not until almost half-a-million years later, with the arrival of Australopithecus afarensis, that we find a truly habitual bipedal hominin.