Morphological and mtDNA analysis of Mezzena mandible

 

I've written about late Neandertals becoming more AMH-like, and a new study on an Italian specimen that postdates the arrival of AMH in Europe lends some further support to that idea.
The Mezzena jaw has Neandertal mtDNA and shares a number of morphological traits with other Neandertals, but its overall shape places it within the cluster of modern humans  (triangles; figure on the left). Notice also that Qafzeh 9 (Q9) and Shkul V (SV) are also within the cluster of modern humans, and Spy 1 (a Neandertal) is actually closer to modern humans than to other Neandertals.
From the paper:

The position on the scatter plot of our specimen of interest, Mezzena, has been calculated a posteriori. Unsurprisingly, the Mezzena mandible does not present any particular affinities with mid-Pleistocene specimens. It is most similar to AMHs being positioned within the H. sapiens cloud of points and the DFA classifies the specimen with modern humans (Table S7). Especially its shape is similar to that of Ohalo II and to a lesser extent to the recent modern human specimen China5. However, it should be noted that its position also indicates affinities with some Neanderthal specimens: the late Neanderthal Spy 1 and Saint-Césaire, the Near-East specimens Tabūn II and Amud 1, and to a lesser extent the classic Neanderthals La Ferrassie 1 and Guattari III (Figure 2).

According to the authors:

In this light, we can interpret the position of the Mezzena mandible which stands within the modern human shape space, while presenting strong shape similarities with some Neanderthal specimens. Such a conflicting taxonomical position is not surprising, considering the geological age of the mandible [30]. Indeed, numerous late Neanderthals such as Spy 1, Saint Césaire and the Near-East mandibles Amud 1 and Tabun II possess hints of a chin (i.e. tuber symphyseo) though not a true modern human morphology [37], [51]. Late Neanderthals lived in area where AMHs might have been already present [2], [23], [52], while the Levantine fossils are displaying a less derived Neanderthal morphology [35], [36]. 

Therefore, in our view, this change in morphology of the mandibular chin among the fossils of Mezzena and other late Neanderthals could have been the result of a small degree of interbreeding with AMHs.

It would be interesting to sequence Mezzena to confirm the existence of AMH admixture.
PLoS ONE 8(3): e59781. doi:10.1371/journal.pone.0059781
Possible Interbreeding in Late Italian Neanderthals? New Data from the Mezzena Jaw (Monti Lessini, Verona, Italy)
Silvana Condemi et al.
In this article we examine the mandible of Riparo Mezzena a Middle Paleolithic rockshelter in the Monti Lessini (NE Italy, Verona) found in 1957 in association with Charentian Mousterian lithic assemblages. Mitochondrial DNA analysis performed on this jaw and on other cranial fragments found at the same stratigraphic level has led to the identification of the only genetically typed Neanderthal of the Italian peninsula and has confirmed through direct dating that it belongs to a late Neanderthal. Our aim here is to re-evaluate the taxonomic affinities of the Mezzena mandible in a wide comparative framework using both comparative morphology and geometric morphometrics. The comparative sample includes mid-Pleistocene fossils, Neanderthals and anatomically modern humans. This study of the Mezzena jaw shows that the chin region is similar to that of other late Neanderthals which display a much more modern morphology with an incipient mental trigone (e.g. Spy 1, La Ferrassie, Saint-Césaire). In our view, this change in morphology among late Neanderthals supports the hypothesis of anatomical change of late Neanderthals and the hypothesis of a certain degree of interbreeding with AMHs that, as the dating shows, was already present in the European territory. Our observations on the chin of the Mezzena mandible lead us to support a non abrupt phylogenetic transition for this period in Europe.
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Etruscan mtDNA origins (Ghirotto et al. 2013)

Etruscan mtDNA origins (Ghirotto et al. 2013)

From the paper:

A model of genealogical continuity across 2,500 years thus proved to best fit the observed data for Volterra, and especially Casentino, but not for another community dwelling in an area also rich with Etruscan archaeological remains (Murlo), nor (as expected) for the bulk of the current Tuscan population, here represented by a forensic sample of the inhabitants of Florence.

and:

As for the second question, the IM analysis shows that indeed there might have been a genealogical link between modern Tuscans and the inhabitants of what Herodotus considered the Etruscans’ homeland, Western Anatolia. However, even under the unrealistic assumption of complete reciprocal isolation for millennia, the likely separation of the Tuscan and Anatolian gene pools must be placed long before the onset of the Etruscan culture, at least in Neolithic times; if isolation was incomplete, the estimated separation must be placed further back in time. Consistent with this view is the observation that Etruscan and Neolithic mtDNAs are close to each other in the two-dimensional plot of Figure S4C; however, a formal test would be necessary to draw firm conclusions from the simple observation of a genetic similarity. Separation times were very close when estimated both using a sample from Western Anatolia, and an expanded sample including individuals from much of Anatolia, and so the choice of the Anatolian population does not seem to affect the results of this analysis.

As always with estimates in years, the choice of mutation rate may affect results, but I am reasonably confident that this particular result does not depend on such issues. From the paper:

For these tests we chose the mutation rate (μ) estimated from the data in the previous ABC analyses (very close to the figure accounting for the time-dependency of the mitochondrial molecular clock [13], μ = 0.003). Tests were also run using the value incorporating a correction for the effects of purifying selection [23] (μ = 0.0014), always finding that it results in a further increase of the estimated separation times (Figure S7B). Only assuming very high mutation rates, at least twice as large as estimated in Henn et al. [13], was it possible to obtain separation times less than 5,000 years (Figure S7B). With both Anatolian samples, any degree of gene flow after separation between the ancestors of Tuscans and Anatolians resulted in more remote separation times.

A couple of observations:

If Etruscans did originate in Anatolia then presumably the historical Etruscans were not descended entirely from them but from a mixture of pre-Etruscans with the incoming population. So, it would seem that the inferred dates are incompatible with a folk migration model of Etruscan origins, but not necessarily with a model that accommodates admixture (e.g., initial mtDNA gene pool separation c. 8,000 years ago with the onset of the Neolithic + later admixture during the Bronze Age). On the other hand, the close similarity between Etruscan and Central European Neolithic mtDNA is a good argument for (mostly) continuity in this case.

That things did happen in Italy in the last 5,000 years can be inferred on the basis of the Iceman's genome. It will certainly be interesting to extract Y chromosomes and/or autosomal DNA from some of these Etruscan samples.

A different issue that may bias dates upwards is the occurrence of East Eurasian mtDNA in current Anatolian Turks. It is not clear by how much this would affect age estimates (this admixture is low, sub-10%, but from a population that split off from West Eurasians perhaps more than 40kya); it would nonetheless be useful to repeat the experiment after either (i) purging the Anatolian sample of lineages likely to have introgressed into the population in medieval times, or (ii) using a different West Asian sample other than that of Anatolian Turks.

In any case, it's great to finally have the genetic characterization of a historical European people, and hopefully more samples will follow both from Italy (at least from those who practiced inhumation) and elsewhere.

PLoS ONE 8(2): e55519. doi:10.1371/journal.pone.0055519

Origins and Evolution of the Etruscans’ mtDNA

Silvia Ghirotto et al.

The Etruscan culture is documented in Central Italy (current Tuscany and Northern Latium, formerly known as Etruria) between the 8th and the 1st century BC. Questions about the Etruscans’ origins and fate have been around for millennia. Herodotus and Livy regarded them as immigrants, respectively from Lydia, i.e. Western Anatolia, or from North of the Alps, whereas for Dionysius of Halicarnassus they were an autochthonous population [1]. Previous DNA studies, far from settling the issue, have raised further questions. The Etruscans’ mitochondrial DNAs (mtDNAs) appear similar, but seldom identical, to those currently observed in Tuscany [2], [3]. Assuming reasonable effects of genetic drift and mutation, these levels of resemblance proved incompatible with the notion that modern Tuscans are descended from Etruscan ancestors [4], [5]. Explanations for this result include the (extreme) possibility that the Etruscans became extinct, but also that their modern descendants are few and geographically dispersed, or that the ancient sample studied represents a small social elite rather than the entire population [4]. As for the Etruscans’ origins, ancient DNA is of little use, because pre-Etruscan dwellers of Central Italy, of the Villanovan culture, cremated their dead [1], and hence their genetic features are unknown. DNAs from modern humans and cattle in Tuscany show affinities with Near Eastern DNAs, which was interpreted as supporting Herodotus’ narrative [2], [6], but in these studies modern Tuscans were assumed to be descended from Etruscan ancestors, in contrast with ancient DNA evidence [5]. The claim that systematic errors in the Etruscan DNA sequences led to flawed genealogical inference [2], [7] is not supported by careful reanalysis of the data [8].

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Redating of the Early Upper Paleolithic site of Riparo Mochi (Italy)

Redating of the Early Upper Paleolithic site of Riparo Mochi (Italy)

There are two possibilities on how the early Aurignacian entered Europe. According to one hypothesis, its bearers followed the Danube, which formed a natural corridor into the heartland of the continent which was, at the time, thickly forested. A different hypothesis is that the early Aurignacian entered Europe via the Mediterranean. Distinguishing between the two hypotheses depends on obtaining reliable chronological estimates for the Mediterranean and Central European Aurignacian

A recent dating of a site in the Swabian Jura suggested that the Aurignacian was earlier attested in Central Europe. But, another paper in the Journal of Human Evolution examines meticulously the sequence in the Moch rockshelter and finds that it is just as early.

Comparisons with dates for other Upper Palaeolithic contexts outside Italy suggest that the date of the Protoaurignacian of Mochi compares closely. In Fig. 9a the start boundaries for the earliest Aurignacian evidence at the sites of Geissenklösterle (Germany), Abri Pataud and Isturitz (France) are compared to the start boundary for unit G in Mochi. The first two sites were dated recently in Oxford with reliable methodologies (Higham et al., 2011; Higham et al., in press) while for Isturitz only a small number of dates exist for the earliest Upper Palaeolithic (Szmidt et al., 2010). This comparison reveals that the lowermost Aurignacian levels at Geissenklosterle (AHIII) and Isturitz (C4d) date to the same period as Mochi G, at around 42.7-41.5 ka cal BP (68.2%). The earliest Aurignacian of Abri Pataud dates slightly later to around 41e40 ka cal BP (68.2%), but the assemblage there has always been considered more evolved, so this is not surprising. No Mousterian dates are included in any of these calculations, therefore the start boundaries in the Bayesian models are not well constrained at their earliest end. What is interesting is that there appears to be a close similarity between the dates for the Protoaurignacian and Early Aurignacian sites in Germany on the Danube and on the Mediterranean coast. This might suggest a rapid dispersal of both variants of the Aurignacian across Europe at c. 44-42 ka cal BP.

It does appear that the Aurignacian was a continent-wide punctuational event in Europe which occurred in the middle to late 40 thousands ka cal BP.

Either there were two streams into Europe (Danubian and Mediterranean), or one stream that quickly inundated much of the continent. Given that the argument for the Danubian Corridor is partly related to the ease of access it provided, it is difficult to imagine how the people who followed it would quickly stray far from it all the way to Italy. Overall, it does appear that there were multiple streams into Europe, and perhaps new research in the Balkans, Eastern Europe, and West Asia, may help us trace the earlier predecssor of these streams before they followed their separate ways into Europe.

Journal of Human Evolution DOI:10.1016/j.jhevol.2011.11.009

A new chronostratigraphic framework for the Upper Palaeolithic of Riparo Mochi (Italy)


Katerina Douka et al.


The rockshelter of Mochi, on the Ligurian coast of Italy, is often used as a reference point in the formation of hypotheses concerning the arrival of the Aurigancian in Mediterranean Europe. Yet, the site is poorly known. Here, we describe the stratigraphic sequence based on new field observations and present 15 radiocarbon determinations from the Middle Palaeolithic (late Mousterian) and Early Upper Palaeolithic (Aurignacian and Gravettian) levels. The majority of dates were produced on humanly modified material, specifically marine shell beads, which comprise some of the oldest directly-dated personal ornaments in Europe. The radiocarbon results are incorporated into a Bayesian statistical model to build a new chronological framework for this key Palaeolithic site. A tentative correlation of the stratigraphy to palaeoclimatic records is also attempted.

Link

Earliest sapiens remains in Europe

Earliest sapiens remains in Europe

From the BBC:

They may be yellowed and worn but these ancient teeth and jaw fragment have something very revealing to say about how modern humans conquered the globe.

The specimens, unearthed in Italy and the UK, have just been confirmed as the earliest known remains of Homo sapiens in Europe.

Careful dating suggests they are more than 41,000 years old, and perhaps as much as 45,000 years old in the case of the Italian "baby teeth".
...

The re-assessments have further importance because palaeoanthropologists can now put modern humans in the caves at the same time as the stone and bone tool technologies discovered there.

There has been some doubt over who created the so-called Aurignacian artefacts at Kents Cavern and the slightly older Uluzzian technologies at Grotta del Cavallo. It could have been Neanderthals, but there is now an obvious association in time with Homo sapiens.

From the NY Times:

They had in fact discovered the oldest known skeletal remains of anatomically modern humans in the whole of Europe, two international research teams reported Wednesday.

The scientists who made the discovery and others who study human origins say they expect the findings to reignite debate over the relative capabilities of the immigrant modern humans and the indigenous Neanderthals, their closest hominid relatives; the extent of their interactions; and perhaps the reasons behind the Neanderthal extinction. The findings have already prompted speculation that the Homo sapiens migrations into Europe may have come in at least two separate waves, rather than just one.

I'll add the abstracts later. This certainly seems to be incompatible with substantial Neandertal interbreeding. If humans and Neandertals were genetically compatible species, then why would they maintain very separate morphological populations for ~10ka? We would expect the two populations to quickly merge into one. Moreover, a longer period of interbreeding in West Eurasia would have left an excess of "Neandertal" ancestry in modern West Eurasians, something we simply don't observe.

Press release:

"What the new dates mean", Benazzi summarised, "is that these two teeth from Grotta del Cavallo represent the oldest European modern human fossils currently known. This find confirms that the arrival of our species on the continent – and thus the period of coexistence with Neanderthals – was several thousand years longer than previously thought. Based on this fossil evidence, we have confirmed that modern humans and not Neanderthals are the makers of the Uluzzian culture. This has important implications to our understanding of the development of 'fully modern' human behaviour. Whether the colonisation of the continent occurred in one or more waves of expansion and which routes were followed is still to be established."

Nature (2011) doi:10.1038/nature10484

The earliest evidence for anatomically modern humans in northwestern Europe

Tom Higham et al.

The earliest anatomically modern humans in Europe are thought to have appeared around 43,000-42,000 calendar years before present (43-42 kyr cal BP), by association with Aurignacian sites and lithic assemblages assumed to have been made by modern humans rather than by Neanderthals. However, the actual physical evidence for modern humans is extremely rare, and direct dates reach no farther back than about 41-39 kyr cal BP, leaving a gap. Here we show, using stratigraphic, chronological and archaeological data, that a fragment of human maxilla from the Kent’s Cavern site, UK, dates to the earlier period. The maxilla (KC4), which was excavated in 1927, was initially diagnosed as Upper Palaeolithic modern human1. In 1989, it was directly radiocarbon dated by accelerator mass spectrometry to 36.4-34.7 kyr cal BP2. Using a Bayesian analysis of new ultrafiltered bone collagen dates in an ordered stratigraphic sequence at the site, we show that this date is a considerable underestimate. Instead, KC4 dates to 44.2-41.5 kyr cal BP. This makes it older than any other equivalently dated modern human specimen and directly contemporary with the latest European Neanderthals, thus making its taxonomic attribution crucial. We also show that in 13 dental traits KC4 possesses modern human rather than Neanderthal characteristics; three other traits show Neanderthal affinities and a further seven are ambiguous. KC4 therefore represents the oldest known anatomically modern human fossil in northwestern Europe, fills a key gap between the earliest dated Aurignacian remains and the earliest human skeletal remains, and demonstrates the wide and rapid dispersal of early modern humans across Europe more than 40 kyr ago.

Link

Nature (2011) doi:10.1038/nature10617

Early dispersal of modern humans in Europe and implications for Neanderthal behaviour

Stefano Benazzi et al.


The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several transitional technocomplexes. Two of these, the Uluzzian of southern Europe and the Châtelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction1, 2, 3, 4, 5, 6. However, the actual fossil evidence associated with these assemblages is scant and fragmentary7, 8, 9, 10, and recent work has questioned the attribution of the Châtelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis11, 12. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal13, 14. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ~45,000-43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals.

Link