Europe

Mitochondrial phylogeography is a useful tool for the study of wild populations. But applying phylogeography to domestic species is more complicated....

A classic example of the use of mitochondrial DNA (mtDNA) diversity to infer the history of domestication refers to dogs (Canis familiaris). Four to six mitochondrial haplogroups (Hg) have been described in genetic studies of modern dogs, indicating recurrent domestication or backcrosses between domestic dogs and wild wolves (Canis lupus). Three of the major Hgs are distributed throughout the world, whereas one (D) is restricted to Europe, especially in breeds originating in Scandinavia. Similar patterns of fragmented genetic diversity have been used to argue for local domestication in other species. Such scenario could apply to dogs as they appear as early as 9000 years ago in Scandinavia, and as dogs and wolf remains have been found on the same sites (Malmström et al. 2008:4).

So, they sampled ancient DNA from Neolithic and medieval dog skeletons, to look for the D haplogroup, which would provide evidence that these ancient dogs had a unique and separate origin from other domesticated dogs elsewhere in the world.

Except, it wasn't there.

Our results indicate that Hg frequencies have been altered in Scandinavian dogs since their first arrival. Interestingly, while Hg C is overrepresented in our ancient material, there is a complete lack of the Scandinavian group D in our ancient dataset. Hg D is the one that could support a Scandinavian origin whereas Hg C is suggested to be of Asian origin. Thus, we find no obvious evidence for prehistoric canid domestication in Scandinavia. An external origin of Scandinavian dogs is supported by morphologic data, as even the oldest remains of dogs in Scandinavia were of smaller size than those of prehistoric and extant wolves. While canid domestication may have occurred in other parts of Europe, Scandinavian dogs were likely imported and had experienced a long period of morphological change under human control before they reached the Scandinavian peninsula (Malmström et al. 2008:7).

Fair enough -- the mitochondrial gene pool of Scandinavian dogs has rapidly changed under human influence during the last few thousand years. No word on where the D haplogroup that characterizes today's Scandinavian dogs has come from; whether introgression from local wolves or dogs elsewhere in Europe.

Why does this remind me of human evolution? Well, consider this 2005 paper by Wolfgang Haak and colleagues:

Here we present an analysis of ancient DNA from early European farmers. We successfully extracted and sequenced intact stretches of maternally inherited mitochondrial DNA (mtDNA) from 24 out of 57 Neolithic skeletons from various locations in Germany, Austria, and Hungary. We found that 25% of the Neolithic farmers had one characteristic mtDNA type and that this type formerly was widespread among Neolithic farmers in Central Europe. Europeans today have a 150-times lower frequency (0.2%) of this mtDNA type, revealing that these first Neolithic farmers did not have a strong genetic influence on modern European female lineages.

I discussed this paper when it came out, noting that one explanation for the results is selection, either in favor of the N1a type in Neolithic farmers or against it later. The change in frequency in post-Neolithic Europeans is clearly not consistent with drift. On the basis of other genetic loci, migration from other populations cannot explain the catastrophic decline in frequency of the N1a type, which the ancient DNA data show was widespread across central Europe. So, there would appear to have been local selection against N1a.

The Scandinavian dogs are showing the inverse pattern -- a now-common mtDNA type was formerly not present at a measurable frequency, at least in the available sample of ancient dogs. With dogs, of course, there is every reason to expect selection imposed by humans. The only question is whether the selection was directly on the mtDNA itself, or whether the mtDNA has been carried along fortuitously with other selected genes. The extreme inbreeding under recent intensive breeding would allow either scenario -- unlike in humans, where no extreme inbreeding occurred.

I want to point out the parallels and differences clearly, because I'm writing this week about effective population sizes and inbreeding. There are many geneticists who hold out the possibility of extreme degrees of inbreeding in post-Neolithic humans. Genetic, archaeological and historic data -- not to mention common sense -- weigh against this possibility. However, many prefer to maintain a strict view that natural selection occurs rarely, if ever.

(via Dienekes)

References:

Malmström H, Vilà C, Gilbert MTP, Storå J, Willerslev E, Holmlund G, Gotherstrom A. 2008. Barking up the wrong tree: Modern northern European dogs fail to explain their origin. BMC Evol Biol 8:71. doi:10.1186/1471-2148-8-71

Haak W, Forster P, Bramanti B, Matsumura S, Brandt G, Tänzer M, Villems R, Renfrew C, Gronenborn D, Alt KW, Burger J. 2005. Ancient DNA from the first European farmers in 7500-year-old Neolithic sites. Science 310:1016-1018. doi:10.1126/science.1118725

Dominique Henry-Gambier and colleagues report in the December Journal of Human Evolution on a newly-discovered cave near Vilhounneur, France, with Gravettian-style parietal art and a partial human skeleton:

A remarkable discovery in France raises anew the question of the relationship between parietal art and funerary practices. France is rich in Gravettian decorated caves, but human remains from this period (28-21,000 years BP) are very rare (Henry-Gambier, 2002). Consequently, the discovery of human and hyena remains in a decorated cave system at "Les Garennes" near the village of Vilhonneur (Charente), just 500 m from the well-known cave of Placard (Clottes et al., 1991), is extraordinary. This new find will result in important new data on the biology and behavior of the Gravettians as well as the disappearance of hyenas at the end of the upper Pleistocene. Here we report on the preliminary results of in-situ observations and radiometric dating of the remains (Henry-Gambier et al. 2007:747).

The skeleton was found in a difficult-to-access chamber with parietal art including a hand stencil:

The walls of the deepest (second) chamber are decorated with red dots, black bars, various traces of color, and a well-executed black hand stencil (Fig. 3a). On a flat surface delimited by concretions, black lines evoke a face (Fig. 3b). Nearby, the partial skeleton (ribs, vertebrae, sacrum, left and right os coxae, left and right femora and tibiae) of a young adult male human is dispersed across the surface of a limestone scree (Fig. 2b). The cranium (Fig. 4) is in a small, low gallery opened just below a painting.

The radiometric date for the skeleton is around 27,000 BP. A number of hyena skeletons from another part of the cave are around 1500 years older, and "were members of the last surviving Pleistocene populations." The authors do not take a position as to whether the human skeleton was a deliberate interment, but compare it to a similar instance from another site:

[T]his site brings to mind the decorated Gravettian cave of Cussac (Dordogne) where several human individuals were placed in bear wallows (Aujoulat et al., 2002).

References:

Henry-Gambier D, Beauval C, Airvaux J, Aujoulat N, Baratin JF, Buisson-Catil J. 2007. New hominid remains associated with Gravettian parietal art (Les Garennes, Vilhonneur, France). J Hum Evol 53:747-750. doi:10.1016/j.jhevol.2007.07.003

National Geographic News' most popular story today is "Odd skull boosts human, Neandertal interbreeding theory."

The NGN article is about a paper coming out in Current Anthropology this month by Andrei Soficaru and colleagues, describing a skull from Pestera Cioclovina, Romania. The skull is between 28,000 and 29,000 radiocarbon years old, and the authors argue that its occipital bone preserves Neandertal-like morphology. The NGN article has some trouble describing the situation, settling for this:

The otherwise human skull has a groove at the base of the back of the skull, just above the neck muscle, that is ubiquitous in Neandertal specimens but has never been seen in the remains of a modern human, argues study leader Erik Trinkaus, an anthropologist at Washington University in St. Louis, Missouri.

"I was frankly quite surprised to see it when I was looking at the specimen," Trinkaus said. "My first reaction was, that shouldn't be there."

The "groove" is a suprainiac fossa, which I can understand is technical, but it's too easy to dismiss things as "a bump here and a groove there" if you ignore the pattern that emerges from which bumps and grooves are there.

Anyway, more on early humans in Europe later. The article ends with an interview with Eric Delson, who is not dismissive but not convinced, either. The final paragraph has this priceless quote:

"But the genetic evidence is not in favor of hybridization, and this fossil does not convince me, nor do the several from Central Europe. I am still waiting for a 'smoking gun,' or perhaps in this case 'a bleeding hand axe.'"

Hmmm....that seems a little like demanding a sign from beyond. I grant, a suprainiac fossa is not exactly stigmata, but hey, a bump here and groove there, and pretty soon you're talking real interbreeding!

The bleeding handaxe. Original photo thanks to Wessex Archaeology, Creative Commons license

I thought I'd link to this article from This Is South Devon. There aren't any real new details, but it sounds like there may be a report on Kent's Cavern soon:

KENTS CAVERN JAWBONE COULD BE EVEN MORE SIGNIFICANT THAN FIRST THOUGHT

Results of tests on a jawbone excavated from Torquay's Kents Cavern are being eagerly awaited to see if the piece is Britain's first example of Neanderthal remains.

The piece has been analysed by the University of Hull's Centre for Medical Engineering and Technology and all that is awaited now is the findings of a detailed CT scan.

The possibility they're hyping is that KC 4 might be a Neandertal:

The research was initiated when Dr Roger Jacobi and Prof Chris Stringer of the Natural History Museum obtained new radio-carbon dates for animal bones found in cave sediments directly above and below where the jaw fragment was found at Kents Cavern.

These indicated that the layer in which the maxilla was found dates to between 37,000 and 40,000 years ago, and, if the jawbone fragment is a similar age, it would be even more significant than first thought.

If the jaw proves to be Neanderthal, then Kents Cavern will not only be the only place in Britain where there is direct evidence that Neanderthals once lived, but also it would confirm that Neanderthals spread across Europe and reached Britain far earlier than is currently thought.

I wrote about the reanalysis of KC 4 in early 2005, and added a post with Keith's diagnosis of the specimen.

I guess if the date is actually 37,000 or earlier, you have to lean toward Neandertal. The specimen is nondiagnostic, and that date would make it earlier than the current earliest modern Europeans (who are from Romania, a lot farther east than Britain). And metrically it is within the range of Neandertals, as I mentioned:

The teeth are highly worn, and their mesiodistal measurements are therefore suspect due to interproximal wear. Even so, they are not outside the range of other Neandertal specimens. The more accurate buccolingual measurements are at the small end of the Neandertal range but not outside it; two specimens from Hortus match the canine and premolar measurements, as does Saccopastore 2. The M1 measurements are the same as those for Spy 1; the B-L breadth of 11.6 is typical for later Neandertals, matching or exceeding specimens from Hortus, Arcy-sur-Cure, Spy, Engis, and La Quina. The molar is not taurodont; and considering that all the teeth are worn essentially flat without occlusal relief, there is unlikely to be any morphological diagnosis based on the dentition.

Scanning is fun and all, but I really doubt that an internal scan is going to reveal anything diagnostic about the specimen (i.e., outside the range of one or the other possibility). And based on the last couple of years of papers, I would say that finding a modern mtDNA sequence would essentially be a negative result: nobody seems to be willing to say that the presence of a modern sequence can be distinguished from contamination.

So, I suppose it will be a Neandertal -- the first known from Britain. I hope the Torquay Museum puts on an exhibit about how hard it is to tell Neandertals from modern humans -- that would be interesting!

I'm just finished with the Neandertal meeting in San Diego, so it's time to decompress a bit. And what better way to do it than some more Neandertal blogging!

It is worth mentioning the paper that finally came out this week about Pestera Muierii, Romania, by Andrei Soficaru, Adrian Dobos, and Erik Trinkaus. Here's the abstract:

The early modern human remains from the Petera Muierii, Romania have been directly dated to 30,000 radiocarbon years before present (30 ka 14C BP) (35 ka cal BP) ("calendrical" age; based on CalPal 2005) and augment a small sample of securely dated, European, pre-28 ka 14C BP (32.5 ka cal BP) modern human remains. The Muierii fossils exhibit a suite of derived modern human features, including reduced maxillae with pronounced canine fossae, a narrow nasal aperture, small superciliary arches, an arched parietal curve, zygomatic arch above the auditory porous, laterally bulbous mastoid processes, narrow mandibular corpus, reduced anterior dentition, ventral-to-bisulcate scapular axillary border, and planoconcave tibial and fibular diaphyseal surfaces. However, these traits co-occur with contextually archaic and/or Neandertal features, including a moderately low frontal arc, a large occipital bun, a high coronoid process and asymmetrical mandibular notch, a more medial mandibular notch crest to condylar position, and a narrow scapular glenoid fossa. As with other European early modern humans, the mosaic of modern human and archaic/Neandertal features, relative to their potential Middle Paleolithic ancestral populations, indicates considerable Neandertal/modern human admixture. Moreover, the narrow scapular glenoid fossa suggests habitual movements at variance with the associated projectile technology. The reproductive and scapulohumeral functional inferences emphasize the subtle natures of behavioral contrasts between Neandertals and these early modern Europeans. (Soficaru et al. 2006:17196).

The paper describes the provenience of the bones -- they are not newly found, but had been originally assumed to be Holocene in age. Recent radiocarbon dating placed them at around 30,000 years old, which makes them among the earliest modern Europeans.

The bottom line is that the bones are modern (i.e., not Neandertal), but they include features that are common in Neandertals. Almost all the other European bones of early Upper Paleolithic date also have Neandertal features. The number and frequency of such features in this earliest Upper Paleolithic sample are greater than in any later sample.

In other words, they look like they have genes from Neandertals. And those genes declined in frequency or effect over time.

Of course for any particular feature on any particular specimen, the story gets more complicated. Take the occipital bun on Muierii 1. It clearly is a projection of the posterior cranium, in the position of the occipital bun in Neandertals, it projects well posterior to inion and it has a fairly abrupt superior aspect. On the other hand, the projection is expressed on a much higher and shorter vault, and certainly doesn't look identical to a Neandertal bun.

But then, Neandertal buns are quite variable, which several Neandertals having no bun at all, and others exhibiting a variable morphology. The ontogeny of the trait probably relates to growth of the posterior brain, the timing of closure of the lambdoidal suture, and the relative bone growth rates of the parietal and occipital bones. Those developmental prerequisites almost certainly differed between skulls with a Neandertal-like cranial shape, and those with a higher, more rounded skull. So the same feature -- or at least, a result of the same developmental process -- may be manifested with different forms in different cranial contexts.

"Contextually archaic" is a nice phrase. It is describing anatomies that occur within modern humans, and that continue to occur within recent and (presumably) living people, but that have become very uncommon. They are far more common in archaic humans, but may have a slightly different pattern of expression, in many cases because the developmental process that generates such features depends on anatomical configurations or events that have themselves changed. So within the context of the sample, they are "archaic" -- reflections back upon earlier humans, in this case Neandertals.

"Neandertal features" certainly has a more intuitive meaning -- features that occur at their highest frequencies in Neandertals -- but it really doesn't convey a lot more information, except for the regional specificity of Neandertals versus all archaic humans elsewhere in the world. But of course since we have many more Neandertals than any other archaic specimens, these "Neandertal features" in some cases are simply "contextually archaic" features in the European context.

What is the point I am coming to? Many "Neandertal features" clearly are more common in early Upper Paleolithic people than in later Europeans, and they show a unidirectional trend toward lower frequencies over time. Some folks would argue that these features don't really demonstrate Neandertal-modern intermixture, because (a) you can't really prove that they are absent in archaic Africans, or they may even be there, although in lower frequencies than Europe; or (b) they are not really the same feature, but instead are consequences of different developmental processes or parallelism.

Why do I think these critiques have little force? Because at this point, we have enough early Upper Paleolithic specimens with such features to notice something very important about them: different specimens have different Neandertal features.

It's like a shotgun approach to Neandertal intermixture. These are not one or two things appearing in parallel, and they aren't chance resemblances in this small early Upper Paleolithic sample, when they almost all decline systematically in later samples.

So when we see each new specimen, like Muierii 1, carrying not only Neandertal features, but its own distinctive set of Neandertal features, that emphasizes the early role of genome-wide intermixture.

However, these traits co-occur with contextually archaic and/or Neandertal features, including a moderately low frontal arc, a large occipital bun, a high coronoid process and asymmetrical mandibular notch, a more medial mandibular notch crest to condylar position, and a narrow scapular glenoid fossa.

Each of these features occurs in other modern specimens, but not in the same combination. And every other specimen from the early Upper Paleolithic with Neandertal or archaic features has a different mix of them. If this phenomenon were the result of parallelism on modern humans entering Europe, or if it were a consequence of features retained from archaic Africans, we should not see this broad and altering mix of features in different specimens.

So bigger samples, adding specimens one at a time, really are important. They let us look at the pattern of variation in ways that test these evolutionary hypotheses.

References:

Soficaru A, Dobos A, Trinkaus E. 2006. Early modern humans from the Pestera Muierii, Baia de Fier, Romania. Proc Nat Acad Sci, USA 103:17196-17201. DOI link

The AP is reporting on the discovery of a double newborn burial near Krems, Austria. The remains are estimated at 27,000 years old, and were buried directly side-by-side along with a string of 31 beads and mammoth bones.

Not much detail in the story, although there is this:

Archaeologists are combing the area to see if the infants' mother is nearby, as giving birth to twins in that era would have been extremely difficult and potentially fatal.

Not to mention for the babies!

This article is about nine months old now, but one of my students brought me a clipping, so I thought I would pass it along. It is a good story about the Pestera cu Oase discovery.

At the start of each day's nine-hour excursion underground, team members stepped into a frigid mountain river that flows into a cave, their helmet-mounted lights piercing the perpetual fog of the cave's 100 percent humidity. As the equipment-laden crew sloshed past stalagmites, the cave narrowed and the air temperature plunged from the 90s to the upper 40s Fahrenheit.

Further in, the ceiling lowered until they were forced, first, to swim on their backs and, finally, don their diving masks and enter a narrow, 80-foot-long underwater passage called "the sump." Underwater visibility was about three feet.

Nice work on why the fossils are important:

Trinkaus made a CT scan of the face to measure the unerupted teeth. "To find wisdom teeth that big," he said, "you have to go back 500,000 years."

In the fair-and-balanced section, the article quotes Richard Klein:

"There could have been interbreeding," Klein conceded. "But all the genetic evidence we have suggests that, if it occurred, it was remarkably rare."

This probably signals more about the reporter's choice of what would heighten the controversy than Klein's actual remarks. On the other hand, if you start hearing an archaeologist talk mainly about the genetic evidence, you have to wonder how weak the behavioral evidence has become.

Mladec 1 (left) and 5 (right), lateral view

A new paper in Nature (May 19, 2005) by Eva Wild and colleagues reports new AMS dates from the Mladec hominid sample. This site has been considered to preserve one of the earliest modern human samples in Europe, but its date has been uncertain. A previous attempt to date a layer overlying the hominid sample resulted in a minimum date of 34,000 - 35,000 years ago (Svoboda et al. 2002), but that date has been questioned. The current paper begins with a great introductory paragraph that reviews the problem of early modern humans in Europe:

The Mladec site has significance for both human evolutionary and archaeological issues and the relevance of its remains has increased as a result of the recent dating of the purportedly Aurignacian-age modern human remains from Velika Pecina (Croatia), Hahnofersand (Germany) and Vogelherd (Germany) to the Holocene epoch, the remains from Koneprusy (Czech Republic) to the Magdalenian period, and those from Cro-Magnon (France) and La Rochette (France) to the Gravettian period. The only directly dated European modern human fossils of Aurignacian age are the Pestera cu Oase (Romania) mandible and cranium at ~35,000 ¹⁴C years before present (that is, ~35 ¹⁴C kyr BP), the Kent's Cavern (UK) maxilla at ~31 ¹⁴C kyr BP, the Pestera Muierii (Romania) remains at ~30 ¹⁴C kyr BP, and the Pestera Cioclovina (Romania) cranium at ~29 ¹⁴C kyr BP, none of which has a secure and diagnostic archaeological association. Moreover, at least the Oase fossils overlap in time with late Neanderthals from for example, Vindija (Croatia), which is at present dated to ~29 ¹⁴C kyr BP and Arcy-sur-Cure (France) at ~34 ¹⁴C kyr BP. The assessment of whether the Mladec fossils are indeed Aurignacian in age, and if so, their chronological position within the Aurignacian time span, has become central to understanding early modern humans in Europe (Wild et al. 2005:332, references omitted).

"Secure and diagnostic archaeological association" is the key element here. There were some modern humans in Europe early, although it is not yet clear that they coexisted in any one place with Neandertals. The Oase remains are sufficient to show the early appearance of the modern human anatomical pattern in Eastern Europe; the appearance in central Europe at Mladec is the subject of the present paper. What there isn't -- as yet -- is any evidence for the idea that modern humans spread new Upper Paleolithic industries into and across Europe.

The directly dated specimens include Mladec 1, 2, 8, 9a, and 25c. Mladec 25c is an ulna, the rest of the specimens were dated from teeth. The dates range from a high of 31,500 (for Mladec 9a) to a low of 26,330 (for Mladec 25c), with most of the specimens between 30,000 and 31,500 radiocarbon years. At this date, the Mladec sample is the oldest modern human sample associated with the later Aurignacian.

There is as yet no diagnostic hominid associated with the earliest Aurignacian. At 31,000 years, Mladec falls nearly 10,000 years after the earliest occurrence of "Aurignacian" assemblages, although what constitutes "Aurignacian" differs a lot between different archaeologists. Since "what is Neandertal" differs a lot between paleoanthropologists, I guess it's only fair.

References:

Svoboda JA, van der Plicht J, and Kuzelka V. 2002. Upper Paleolithic and Mesolithic human fossils from Moravia and Bohemia (Czech Republic): some new ¹⁴C dates. Antiquity 76:957-962.

Wild EM, Teschler-Nicola M, Kutschera W, Steier P, Trinkaus E, and Wanek W. 2005. Direct dating of Early Upper Paleolithic human remains from Mladec. Nature 435:332-335. Nature online

Following up on my earlier post on the Kent's Cavern 4 maxilla: although my library doesn't have back issues of the Proceedings of the Torquay Natural History Society, it turns out that the Center for Research Libraries in Chicago does. So I have acquired a copy of Keith's short report on the Kent's Cavern 4 specimen.

The measurements of the teeth reported by Keith are as follows:

The teeth are highly worn, and their mesiodistal measurements are therefore suspect due to interproximal wear. Even so, they are not outside the range of other Neandertal specimens. The more accurate buccolingual measurements are at the small end of the Neandertal range but not outside it; two specimens from Hortus match the canine and premolar measurements, as does Saccopastore 2. The M¹ measurements are the same as those for Spy 1; the B-L breadth of 11.6 is typical for later Neandertals, matching or exceeding specimens from Hortus, Arcy-sur-Cure, Spy, Engis, and La Quina. The molar is not taurodont; and considering that all the teeth are worn essentially flat without occlusal relief, there is unlikely to be any morphological diagnosis based on the dentition.

From the description, it appears that the lateral view of the maxilla reveals nothing diagnostic, either. Keith reports that "there is preserved a small area of the lower wall of the sinus maxillaris, with the basal part of the zygomatic ridge of the upper jaw" (Keith 1927:1). I'm still hoping to find a picture of that side; my copy did not include Keith's figure.

Keith's diagnosis of the specimen is as follows:

Nor can there be any doubt as to the nature of the individual represented by this fragment: the teeth in their dimensions and characteristics agree in every detail with those from jaws of men of the modern type. And in this type I include, of course, the late palaeolithic peoples of Europe. The teeth and jaw now described may very well have belonged to the same people whose remains have been already discovered in Kent's Cavern -- namely the palate found deep in the upper stalagmite by Mr. William Pengelly, and the other specimen found near the mouth of the cavern and described in the last number of this journal. One can say with assurance that the specimen now described could not be derived from an individual of the Neanderthal type. Further, from the dimensions of the teeth I infer that the individual represented by the specimen was of the male sex and the degree of the wear shown by the crowns of the teeth indicate that he had reached middle life (Keith 1927:1-2).

Keith reports that the specimen is modern, but this is of course in the context of 1927, when Keith and many others believed that modern humans had a long antiquity as Neandertal contemporaries. Thus the not-so-subtle triumphalism associated with every find that appeared to place "modern" humans early in the Paleolithic (the first paragraph of this piece goes to great length to argue for the antiquity of the specimen). The question is not so different today, particularly since this would be the earliest modern human specimen in Europe if it is modern and if the 40,000 year date is accurate. But today we have a broader knowledge of the anatomy of late Neandertals, and this specimen appears to fit within that range as well as the range of modern humans.

Will DNA testing settle the issue? I don't really think so. What does it mean if we find a Neandertal sequence? If the specimen is a modern human at the western edge of Europe 40,000 years ago, what modern human would be more likely to have a Neandertal sequence? Without a strong anatomical case, what is to dispute the hypothesis that this specimen belonged to a population with a mixture of Neandertal and modern morphologies? The same questions could be asked if the date turns out not to be accurate, especially if the prior 31,000 year date was the correct one.

References:

Keith A. 1927. Report on a fragment of a human jaw found at a depth of (10 1/2 ft) 3.2 m in the cave earth of the vestibule of Kent's Cavern. Trans Proc Torquay Nat Hist Soc 5: 1-2.

Kent's Cavern, near Torquay, England, underwent systematic archaeological investigation beginning in the 1860's, proceeding intermittently up to the present day. There is a substantial Middle Pleistocene record of human occupation in the cave. The most important fossil human specimen is the Kent's Cavern 4 maxilla, preserving the right canine, third premolar and first molar and the bone holding them together, with a small piece of palate.

Kent's Cavern 4, medial view

According to a story in This Is Devon, new AMS dating by the Oxford Radiocarbon Accelerator Unit has placed the maxilla between 37,000 and 40,000 years ago.

The date should be treated with caution, lying as it does in a time range that is very difficult to date accurately with AMS method. If it is accurate, then the maxilla is older than the Oase fossils from Romania. The key point of interest now is its taxonomy. From the article:

Barry Chandler, assistant curator at the museum, where the jaw bone is currently on display, said the new conclusions posed fresh questions. He said: "If the jaw is anatomically modern - from humans known as Cro-Magnons as Keith believed - then these people spread across Europe, reaching Britain far earlier than is currently thought.

"If, however, Keith was wrong and the jaw is from the human species known as the Neanderthals we will have the first direct evidence of Neanderthals on mainland Britain. We hope to resolve this problem by extracting ancient DNA from one of the teeth."

The Keith (1927) reference is below; I haven't seen it so I don't know what the argument is based upon. Looking at the maxilla, I just don't think there's enough there to make this diagnosis. It should be recalled that Keith's assessment was made during a time when the archaeological sequence was much less established, and there was an active effort to establish the existence of Homo sapiens alongside or previous to the Neandertals. My guess would be that this is a Neandertal; as Churchill and Smith (2000) describe it, there is no compelling association to a specific archaeological industry, so without a strong anatomical case there is no reason to think it is not Neandertal. I have not, however, seen the lateral side of the specimen. If anyone has a lateral or frontal view, I'd be really happy to see it. It could surprise me by being really informative, but even if it was clear whether a canine fossa or maxillary notch may have been present I'm not sure that would be sufficient to prove the specimen is modern.

At the same time, I'm not sure this is the best instance to try taxonomy-by-DNA. In fact, the most interesting possible result (a modern human DNA sequence) would be fundamentally equivocal. Would that mean this was a modern human? Or a Neandertal with a modern human sequence? Or contamination? There is really no way to tell, and that means that only the less interesting result (a sequence clustering with the Feldhofer specimens and other Neandertals) would be informative.

UPDATE: A new BBC story by Paul Rincon gives more details about the dating and potential study of the specimen. The new radiocarbon testing was performed because the old 31,000-year date may have been contaminated by glue applied to the specimen after excavation. There is also this about the Neandertal/modern human assignment:

Further research on the jawbone fragment is planned with the aim of answering this question.

Chris Stringer, of London's Natural History Museum, and Erik Trinkaus, of Washington University in St Louis, US, will carry out a physical examination of the specimen to see if it carries any features diagnostic of either modern humans or Neanderthals (Homo neanderthalensis), their close cousins.

My only concern in this whole thing is the evident unwillingness of anybody to just look at the thing sitting on the museum shelf, until the current redating made it an issue. This is not a snipe against Stringer and Trinkaus at all -- first, because I can think of few better to settle this question, and second because they usually have better things to do than look at every bit of maxilla (albeit one of the only human remains associated with the early Upper Paleolithic in western Europe!). But how many clever British graduate students have there been who could have hopped a train to Torquay and done this observation 10 years ago? 25 years ago?

Let's hope this will make some graduate students consider where they might find the next Kent's Cavern 4.

The Palanth forum has spawned a nice thread on the Kent's Cavern dating, with references and additional information.

If you are interested in the history of the Kent's Cavern excavations, or even in visiting the place itself, there is a very nice virtual tour hosted by the proprietors.

References:

Churchill SE, Smith FH. 2000. Makers of the Early Aurignacian of Europe. Yearbk Phys Anthropol 43:61-115. Wiley InterScience

Keith A. 1927. Report on a fragment of a human jaw found at a depth of (10 1/2 ft) 3.2 m in the cave earth of the vestibule of Kent's Cavern. Trans Proc Torquay Nat Hist Soc 5: 1-2.

Mellars P. 2005. The impossible coincidence. A single-species model for the origins of modern human behavior in Europe. Evol Anthropol 14:12-27.

I happened across an article by Pavlov and colleagues (2001) about the Mamontovaya Kurya site in the Russian Arctic. From the abstract (64):

The transition from the Middle to the Upper Palaeolithic, approximately 40,000-35,000 radiocarbon years ago, marks a turning point in the history of human evolution in Europe. Many changes in the archaeological and fossil record at this time have been associated with the appearance of anatomically modern humans. Before this transition, the Neanderthals roamed the continent, but their remains have not been found in the northernmost part of Eurasia. It is generally believed that this vast region was not colonized by humans until the final stage of the last Ice Age some 13,000-14,000 years ago. Here we report the discovery of traces of human occupation nearly 40,000 years old at Mamontovaya Kurya, a Palaeolithic site situated in the European part of the Russian Arctic. At this site we have uncovered stone artefacts, animal bones and a mammoth tusk with human-made marks from strata covered by thick Quaternary deposits. This is the oldest documented evidence for human presence at this high latitude; it implies that either the Neanderthals expanded much further north than previously thought or that modern humans were present in the Arctic only a few thousand years after their first appearance in Europe.

An interesting start, that. The archaeology includes a mammoth tusk with marks that may be the result either of deliberate incision or of chopping of other material using the tusk as an anvil. The radiocarbon date of bones and the tusk range between 34,400 and 37,400 years ago.

There are no fossil humans at this site. The authors raise the issue of attribution, noting that the date of the site would mean that modern humans had expanded into the Arctic fringe very shortly after they appeared in Europe.

A pressing question is whether the pioneers who lived in these northern landscapes were members of the ancient Neanderthal population (Homo sapiens neanderthalensis) or newcomers from the south. Most scholars associate the Aurignacian industry -- the more advanced stone-tool technology that appeared in Europe at around 40,000 yr BP -- with the emergence of modern humans. However, the earliest indisputable remains of humans with a fully modern morphology (Homo sapiens sapiens) date to 30,000-35,000 yr BP; that is, well after the archaeologically defined transition from the Middle to the Upper Palaeolithic. In European Russia, well preserved skeletons from the famous Palaeolithic site of Sungir, northeast of Moscow (Fig. 1), show that anatomically modern humans were present there not later than 28,000 yr BP. At the Kostenki IV site on the west bank of the Don river, bones of modern humans have been uncovered from strata dated to 30,000 yr BP. The stone-working technology reflected in the Byzovaya material is similar to that of Sungir and other early Upper Palaeolithic sites of the eastern Szeletien tradition, indicating that these artefacts were manufactured by modern humans. However, whether the person who inflicted the marks on the tusk from Mamontovaya Kurya, as much as 8,000-9,000 years earlier, belonged to the same human lineage as the residents at Byzovaya and other Palaeolithic sites further to the south is more uncertain (Pavlov et al. 2001:66-67, citations omitted).

They also relate the site extensively to the Byzovaya site in the preceding paragraph, noting the archaeology has been classified as "eastern Szeletien with Aurignacian traits" (66).

I remembered Szeletian as a Neandertal-associated "transitional" Upper Paleolithic industry, so I went on a bit of a search to see what the story actually is. The search is not finished yet, as there are some books to consult, but the short answer is that Szeletian means different things to different archaeologists. From Churchill and Smith (2000:75, citations in original):

Most of the various IUP [Initial Upper Paleolithic] cultures of Central and Eastern Europe that are characterized by leaf points have at one time or another been seen as regional variants of the Szeletian, a culture defined at Szeleta Cave in the Bukk Mountains of Hungary (Allsworth-Jones 1990a). Regional distinctions can be identified, however, and the possibility exists as well that the occurrence of leaf points in different regions may be the result, at least in some cases, of convergence (Allsworth-Jones 1990a). Regionally defined leaf-point cultures include the Brynzeny, the Gordineshty, and the Kostenki-Streletsian (Streletskaya) of the Russian Plain (Anikovich 1992), the Altmuhlian of southern Germany, the Jerzmanowician of eastern Germany and Poland, the Bohunician of the Czech Republic (in which leaf points are relatively rare, leading to suggestions that this industry should be considered as a development separate from the Szeletian: Svoboda 1998, 1990), and the Jankovichian (although this might be seen as a Middle Paleolithic industry with leaf points: Gabori-Csank 1990) and the Szeletian sensu stricto of Hungary (see Allsworth-Jones 1990a).

Whoa. At a glance, it sounds like like the Szeletian is defined by leaf points in about the same way that the Aurignacian is defined by split-base bone points, and is therefore just about as unitary -- in other words, it barely hangs together. Like I said, I'm working on understanding this better, but my predilection is to step away from the names and consider the possibility that the "Szeletian" really is a constellation of behaviors weakly if at all linked, and it therefore has no necessary link to biological differences between human populations.

The argument for the Szeletian being the product of the Neandertals is almost entirely based on the similarities between it and the earlier Micoquian industry. There are only two sites with human remains associated with Szeletian assemblages, and neither of these has been clearly shown to be Neandertal in anatomy (the sum total is four teeth from both sites) (Churchill and Smith 2000). The case for archaeological similarity, and the status of the Szeletian as a "transitional" industry, is presented by Valoch (2000):

The situation with the Szeletian is quite different. Even though it is also likely to have been produced by the Neanderthals, the stone industry differes from the Bohunician in completely lacking a Levalloisian component. Typological and technological analyses have shown that the archaic elements of the industry are Micoquian and reflect a technological complex that was widespread in Central Europe. However, the types characteristic of the Upper Palaeolithic are carinated and nosed scrapers and carinated burins -- shapes exclusive to the Aurignacian (Allsworth-Jones 1986, 1990; Oliva 1991, 1992; Valoch et al. 1993). The only type specific to the Szeletian -- the leaf point -- has its origin in the Micoquian. The genesis of such an industry can be explained in only two ways: either it developed as a result of spontaneous substrate evolution (i.e., Micoquian), in which case the different Aurignacian types developed in parallel or almost simultaneously and quite independently of the Aurignacian proper, or the Aurignacian had a share in the formation of the industry through some form of contact with the Micoquian. No other explanation appears viable today, although future studies may produce new information (Valoch 2000:625).

The argument for the eastern Szeletian being the product of modern humans apparently comes from the association with the remains at Sungir. Since these apparently are not necessarily the same cultural tradition as other Szeletian sites (despite the shared name), there seems not to be a conflict.

The implication that the far northern tier of Eurasia was occupied very early by modern humans is another piece of evidence consistent with the idea that the first modern Europeans came from the far north. This hypothesis proposes that the features that people spread into the Palearctic as a rather specialized adaptation, and may have exploited a niche available to highly mobile, long-limbed, and culturally sophisticated people. Ultimately, the eastern extreme of this population may have moved into Beringia and further to the New World.

A list of online resources related to the topic of Paleolithic occupation of the circumarctic is maintained at WorkingDogWeb, which I assume is related because of dogsleds?

References:

Churchill SE, Smith FH. 2000. Makers of the Early Aurignacian of Europe. Yearbk Phys Anthropol 43:61-115. Wiley InterScience

Pavlov P, Svendsen JI, Indrelid S. 2001. Human presence in the European Arctic nearly 40,000 years ago. Nature 413:64-77. Nature

Valoch K. 2000. More on the question of Neanderthal acculturation in Central Europe. Curr Anthropol 41:625.