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nds nature online article doi 10 1038 news041220 13 moore m j early g a 2004 cumulative sperm whale bone damage and the bends science 306 5705 2215 tags zoology comments 2 comments categories anatomy evolution open access osteology osteonecrosis sperm whales zoology anatomically modern humans a brief introduction 22 apr the imperative of the human species to know thyself has developed into a rapidly expanding field in palaeoanthropology the exploration of our species homo sapiens is a particularly active field which utilizes multi disciplinary approaches to untangle the evolutionary threads of our beginning the following essay introduces concepts and approaches used in this field whilst raising current research issues for a species that is both narcissistic and inquisitive homo sapiens has so far done a remarkably poor job of defining itself as a morphological entity tattersall and schwartz 2008 49 thus starts the opening sentence to tattersall and schwartz s 2008 article on the problems of clarifying the morphological distinctiveness of anatomically modern humans amh or the species homo sapiens it is perhaps applicable not just to the morphological characteristics but also the fossil record and origins of amh themselves pearson 2008 38 this paper then will discuss the principles behind the definitions and evolution of amh in context with reference to its behaviour and morphological traits in turn the dominant models of the origin and subsequent dispersion of amh will be discussed with reference to where homo sapiens fit in the palaeoanthropological record a wealth of new genetic research data and fossil finds has considerably opened up the treasure chest of hominin information which is having a considerable impact on our understanding of the h sapiens place in the evolutionary records bowden et al 2012 curnoe et al 2012 krause et al 2010 prat et al 2011 wood 2005 42 it is directly as a result of how the reporting of evolutionary science has changed in the past few decades mcewan 2012 and how technological approaches have uncovered so much genetic data in reconstructing fossil record relationships jurmain et al 2011 270 that the definition of amh is not so easy this paper will conclude with a talk on how the biocultural evolution of h sapiens is now impacting both our environment and localised populations in certain contexts le fanu 2009 hawks et al 2007 jurmain et al 2011 it is important to note that h sapiens are the last species of the genus homo with the first species tentatively dated in africa to nearly 2 5 million ya years ago which led to the first dispersal of hominins largely h erectus from africa around 1 8 ya jurmain et a l 2011 240 amh dispersal occurred much later it was once thought that amh were defined by modern anatomy and behaviour at the junction of the upper palaeolithic around 40 000 ya nowell 2010 438 however recent palaeoanthropological finds and research have discovered a distinct decoupling between early amh anatomy and later symbolic modern behaviour with anatomically similar traits of amh in fossils pinpointed to east and south africa to around 200 000 ya rightmire 2008 8 wood 2005 however there are problems concurrent with the dating of the hominin fossil record as millard 2008 870 concludes that the dating evidence for many key fossils is poor typically there are a number of assigned morphological features that mark out homo sapiens compared to other species in the homo genus table 1 as tattersall and schwartz 2008 51 note however impressive the suite of features not all of them are expressed with equal emphasis in all living humans when this is combined with the fossil record of amh with individuals often taken as examples for their own long lost skeletal population and the problems inherent in the preservation of skeletal elements geological pressure scavenging etc we should rightly be wary of definitively assigning a species name before comparison with relative contextual remains stratigraphic layers and other similar period sites millard 2008 pettitt 2005 general characteristic morphological features of amh cranial cranial capacity in excess 1350cc variable distinct chin inverted t relatively veretical frontal bone relatvely flat non projecting face brow ridge expressed more clearly in males round occipital region small incisor teeth post cranial narrow thorax small and narrow pelvis straight limb bones typically less robust more gracile then recent ancestors table 1 general morphology for homo sapiens pettitt 2005 132 tattersall and schwartz 2008 51 wood 2005 110 nb see also pearson s table 2 2008 39 using a cladistics framework pearson 2008 38 highlighted the fact that there are specific difficulties in using statistical measurements of metrical and discrete measurements as having been conceptualised as derived features in amh crania with comparison to neandertal and h erectus crania however there are further problems when trying to establish if the earliest h sapiens african fossils of omo kibish the herto crania or near eastern skhul and qafzeh fossils fit within the 95 rate of modern features with results not even reaching the 75 fit of the modern features for amh pearson 2008 39 in part this is due to fossils such as the herto crania which are used as the mean of that particular population which ultimately conflates individual within population variation and between population variation jurmain et al 2011 pearson 2008 39 other problems of quantifying such long chronological morphological differences include the lack of various populations of modern australian aboriginals for example and certain prehistoric peoples being outside of the 95 confidence to fit the given morphological concept of amh clearly there needs to be a control on the temporal geographic population of the amh under consideration in such studies when carrying out both the statistical analysis with other fossil hominins and when taking the defining measurements pettitt 2005 132 137 argues that h sapiens should be classed into three arbitrary chronological groups of morphological continuity 1 those of the earliest h sapiens including material from bodo ethopia broken hill zambia and elandfontein south africa amongst others 2 transitional or archaic h sapiens including herto omo kibish 1 and 2 ethiopia florisbad south africa and jebel irhoud morocco 3 finally amh including makapansgat border cave and equus cave south africa taramsa egypt and dar es soltan morocco examples see table 2 below for dates this ordering of morphological continuity defines amh through the evolution of h sapien traits with retention of h ergaster traits earliest whilst the amh group compromise clear amh dating to less than 125 000 ya pettitt 2005 132 as pearson 2008 44 suggests the process of becoming modern likely occurred as a series of steps regardless of whether one considers these different steps to be different taxa in a bushy phylogeny or merely different grades in a single evolving lineage pearson 2008 44 goes on to say that the evolution of modern man should be viewed as a process rather than an event involving rapid morphological change due to drift during population bottlenecks and selection for new advantageous traits or genes or a combination of the two rather than a singular smooth process therefore we should be wary of relying purely on the often sparse fossil record regardless it is widely recognised that h sapiens are a probably daughter species of h erectus i e as a result of a speciation occurrence which spread across africa and into western eurasia at the beginning of or just before the middle pleistocene jurmain et al 2011 rightmire 2008 8 recent research has also led to five majority agreements in regards to the tenets of amh behaviour table 2 nowell 2010 447 wood 2005 109 makes the salient point that early eurocentrism in the search for amh behavioural origins clouded certain judgements such as focusing on western europe to the detriment of african archaeological sites points of consensus on modern behaviour the relationship between modern anatomy and modern behaviour is more complex than once thought modern behaviour has symbolic thoughts at its core archaeological record of the african middle stone age has rendered invalid the idea of a human revolution occurring for the first time in the upper palaeolithic of western europe later neandertal sites have demonstrated modern behaviour to either some form or some degree such as personal adornment or symbolic behaviour the triad of social cultural and demographic factors are key in understanding variability and patterning in the archaeology record table 2 agreed points in visioning the concept of modern behaviour balter 2011 21 nowell 2010 447 pettitt 2005 zilhao 2006 2010 1025 research jurmain et al 2011 prat et al 2012 has also highlighted symbolic behaviour in a number of early h sapiens sites throughout africa and the near east balter 2011 21 highlights aterian sites in north africa where various personal and possible symbolic artefacts have been found whilst blombos cave in south africa 77 000 ya and katanda in the dr of congo 80 000 ya have some of the earliest symbolic artefacts recovered including incised ochre worked bone and beads almost a full 45 000 years before any such artefacts appear in the european record jurmain et al 2011 298 299 mellars 2006 9383 proposes a model that indicates climatic environmental and cultural changes around 80 000 to 60 000 ya as major causative agents of cognitive change alongside population pressures in the dispersal of african h sapiens however nowell 2010 441 states that the gradual emergence of behaviours as a mosaic of features and not as a single revolutionary package should be considered within the archaeological record whilst defining that for the majority of researcher s symbolic language and codified social relationships define modern behaviour mosaic features in fossil hominids have been noted in recent discoveries of the australopithecus sediba specimen highlighting a mix of australopithecus and homo anatomical features wong 2012 25 the origins of amh living outside of africa have led to the formation of two major competing models in palaeoanthropolog the multi regional continuity hypothesis that proposes already living populations of hominins and local populations in asia europe and africa continued their indigenous evolutionary development from pre modern middle pleistocene forms to anatomically modern human jurmain et al 2011 281 whilst the complete replacement or out of africa hypothesis proposes that amh arose in africa 200 000 ya to completely replace those in europe and asia table 3 jurmain et al 2011 279 critical to the multi regional hypothesis are the tenets that i a level of gene flow between geographically separated populations prevented speciation ii all living humans derive largely from the species h erectus iii natural selection in regional populations is responsible for the regional variants found in extant populations and finally iv that the emergence of h sapiens was not restricted to one area per se but was a phenomenon that occurred throughout the geographic range where humans lived johanson 2001 1 table 3 timeline of major h sapiens discoveries question marks denote tentative dates jurmain et al 2011 413 click to enlarge critical to the complete replacement theory are that i h sapiens arose in one place highly likely to be east south africa ii h sapiens ultimately migrated out of africa and replaced all human populations without interbreeding and that iii modern human variation is a relatively recent phenomenon johanson 2001 1 although not all factors of the multiregional hypothesis cannot be falsified it seems prevalent that h sapiens originated in eastern africa with ethiopia so far providing the most stable dated site and dispersed to europe and asia from 65 000 ya onwards in various waves table 2 jurmain et al 2011 282 mellars 2006 9381 the two most securely dated sites in europe for amh are pecstera cu oase in romania at 42 000 ya and buran kaya iii in the crimea ukraine at 31 900 ya hoffecker 2009 16040 prat et al 2011 unsurprisingly hoffecker 2009 16040 notes that the issue of the mechanism of transition is a controversial topic in palaeoanthropology arguments have been made that amh crossed into eurasia via a levantine corridor with the earliest amh dates from skhul and qafzeh in israel at around 120 000 to 100 000 ya wood 2005 98 whilst recent work in north african aterian populations from around the same period are pointed out as being possible ancestors to at least some of the h sapiens who left africa during this period balter 2011 23 the palaeoanthropological evidence suggests that they the aterians possessed the right symbolic behaviour anatomy and favourable climatic conditions to be at least a contender for contributing to one of the waves of h sapiens leaving balter 2011 22 23 there are a variety of sites across europe after 40 000 ya that show a variety of evidence for amh presence including the triad of modern human behaviour with symbolic artefacts and modern skeletal morphology however we should not forget that europe was already populated with the h neandertalensis species prior and co existed with h sapiens for approximately 10 000 years or so hoffecker 2009 16040 wood 2005 110 this subject will be tackled shortly the most secured dates found in asia are from areas such as the sahul region conjoined landmass of australia papua new guinea and tasmania where it is possible amh occupied various areas wood 2005 111 112 it must be remembered that while the dwarf species h floresiensis survived up until 18 000 ya on the island of flores with temporal overlap between themselves and h sapiens it seems unlikely there was regional overlap from the archaeological evidence wood 2005 111 curnoe et a l 2012 1 note that the amh fossil record for east asia is at this time poorly recorded owing to a lack of detailed description rigorous taxonomy classification and a distinct lack of accurately dated fossils however there are a few key sites liujiang in southern china has produced a skeleton which although it lacks exact stratigraphic position has been dated to an estimated broad range from 153 30 000 ya whilst the niah cave child in east malaysia has been dated to 45 39 000 ya for the cranium from a recent field and lab program curnoe et al 2012 2 tianyuan cave just south of the zhoukoudian cave has fragmentary evidence of an amh crania and teeth which are dated to 40 000 ya with a possible mix of archaic and modern features the american and chinese team who excavated it have suggested it is evidence of interbreeding in china with resident archaic populations but suggest an african origin for the amh itself jurmain et al 2011 287 the above examples highlight problems in understanding the definition of amh both anatomically and behaviourally with the advent of dispersals from africa am...