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paleoanthropology, genetics and evolution

Chatelperronian

  • Neandertals didn't disappear before 40,000 years ago

    Tue, 2011-05-10 19:25 -- John Hawks

    The science press has its own synchronized cycle, like brain waves, and being in Rome seems to make me into a misfiring neuron. Here it is tomorrow, and there's this story about Neandertals all being dead before modern humans showed up, which for Americans is now yesterday's news. Unless you take the paper NY Times, of course, in which case you probably haven't read it yet.

    The occasion for the article is a paper reporting new radiocarbon dates for one of the specimens from Mezmaiskaya, a site in the Russian Caucasus.

    The site and the date of the Mez 2 burial

    Excavations by Golanova and colleagues have recovered two burials of young children from this site. One of them (Mez 1) has been the subject of much research. Based on some skeletal features and a partial sequence of its genome, the skeleton is a Neandertal child. A sample of one of its ribs was taken for radiocarbon dating, reported in 1999, and yielded a direct AMS date of 29,000 BP. This was out of sync with the other dates from the surrounding level of the site.

    Ten years ago, Milford Wolpoff and I suggested that the skeletal features by themselves weren't very convincing, and a recent date (apparently out-of-sync with the surrounding archaeological layer) might signal an intrusive Upper Paleolithic burial [1], despite its Neandertal mtDNA sequence. Now we can look at a large part of the genome of this individual, which is very much like the Vindija Neandertals. In 2005, Skinner and colleagues reported ESR dates from Mezmaiskaya, concluding that layer 3 (including the Mez 1 burial) dates to between 60,000 and 70,000 years ago. By that time, those authors were discussing the inconsistency of the recent 29,000 date for the rib, compared to much older dates (>35,000 BP) for an overlying Mousterian layer. They expected the underlying layer 3 to be much older, and found that to be true of the ESR date estimates.

    The second child burial, Mez 2, comes from layer 2 of the site, which is also Mousterian but younger than the first burial. A date around 40,000 years for the Mez 2 infant is basically what was expected six years ago by Skinner and colleagues:

    Infant 2 was found in a pit introduced from Layer 2 into Layers 2A and 2B(1). Its’ age therefore is probably about 40 ka. Since the precise surface from which the pit was dug is unknown, this should be considered a maximum age.

    That left open the possibility that the burial might be younger.

    In the new paper [2], Ron Pinhasi and colleagues report that a sample taken from the infant itself dates to 39,700 +/- 1100 radiocarbon years BP, which calibrates to between 42,960 and 44,600 calendar years BP. The new date confirms that the burial happened relatively soon after the deposition of the surrounding dated bones.

    The authors additionally report many other date estimates for faunal materials from the site. These form a pattern in which most are consistent with a relatively narrow range of date estimates, but a few are outliers. One of the important conclusions from the outliers is that contaminated carbon is hard to get out of a sample, even with the advanced ultrafiltration performed by the Oxford lab. The conclusion is narrowly interesting and solid, and it's very important to iron out such inconsistencies -- compare, for example, my 2008 post on the Gorham's Cave chronology.

    So what is the big deal?

    What does the paper say about the dates of other sites?

    Here's where things get interesting. The paper includes this passage in its discussion:

    The critical reanalysis of directly dated Neanderthal and AMH fossils from across Eurasia, taking into consideration pretreatment histories and redating results (5), supports our findings in the Caucasus and highlights the lack of reliably dated Neanderthal fossils younger than ∼40 ka cal BP (Fig. 3). Contrary to traditional arguments for up to 10,000 y of coexistence, these data suggest that Neanderthal extinction across Western Eurasia, including the Caucasus, was probably a rapid process, and that coexistence with AMHs, when it occurred, may have been of limited duration.

    and this in its abstract:

    Our results confirm the lack of reliably dated Neanderthal fossils younger than ∼40 ka cal BP in any other region of Western Eurasia, including the Caucasus.

    That last part is a pretty strong statement. No reliably dated Neandertal fossils anywhere after 40,000 years ago?

    I thought that was so surprising that I corresponded with the study authors today. One distinct advantage of being in Rome is that I'm synchronized with Europeans, so Tom Higham was able to write back with some of his thoughts. The authors' doubt in the later dates for Neandertal specimens is genuine; their experience is that the newer treatments to remove recent contaminating carbon from samples is eliminating Neandertal dates under around 40,000 years.

    A systematic revision of the radiocarbon chronology of late Middle and early Upper Paleolithic Europeans has been underway for several years. This has been an important story, and I've written about it several times (my "dating" category hits most of the posts). I think I once told a journalist that this was the most underreported story in paleoanthropology.

    In 2006, Higham and colleagues reported that dates obtained for the Vindija G1 Neandertals, at 29,000 BP, were too young by some 4000 years [3]. That result is listed in the current paper as "doubtful" becuase it did not employ the latest purification strategies. That helps to show that the current paper is "equal opportunity" -- past results from the Oxford Accelerator unit are not immune to doubt. But it is hardly confidence-raising. If we cannot trust radiocarbon determinations made in the last five years, why should anyone submit further samples for testing?

    Personally, this was my reaction to the paper: don't grind up any more human bone until the radiocarbon community is unified about sample processing techniques. Let them work it out on the fauna.

    The paper lists 15 direct dates on Neandertal specimens younger than 40,000 calendar years BP (some of them multiple samples from single skeletal remains). It lists all 15 of these as doubtful because they do not employ the latest techniques. That is a point emphasized by Higham also (and reflected in several past papers): these date determinatinos are not trustworthy given what we know about sample contamination by recent carbon-14. The Oxford group has put out several papers on this problem. One of the most useful is by Blockley and colleagues [4] because it introduces the device of using the Cantabrian Ignimbrite ash horizon as a marker to compare dates -- dates below the horizon should be consistent, whereas a large sample of actual date estimates include many that are far too young.

    At any rate, this is where the story in the linked news article (and others) comes from. University College Cork issued a press release in conjuction with the paper's early edition release in PNAS.

    Direct dating of a fossil of a Neanderthal infant suggests that Neanderthals probably died out earlier than previously thought. Researchers have dated a Neanderthal fossil discovered in a significant cave site in Russia in the northern Caucasus, and found it to be 10,000 years older than previous research had suggested. This new evidence throws into doubt the theory that Neanderthals and modern humans interacted for thousands of years. Instead, the researchers believe any co-existence between Neanderthals and modern humans is likely to have been much more restricted, perhaps a few hundred years. It could even mean that in some areas Neanderthals had become extinct before anatomically modern humans moved out of Africa.

    This is the lead of the press release. I think that the claim makes up only a minor part of the paper (which is really a results paper about Mezmaiskaya). It is clearly interesting and provocative, but I think the paper's results by themselves do not justify the claim. In the case of Mez 2, a skeleton that the excavators expected to be 40,000 years old, actually turns out to be 40,000 years old. No surprise. There were no incorrect radiocarbon assessments of this specimen, and the apparently wrong assessment of the Mez 1 infant (at 29,000 years old instead of beyond radiocarbon range) has not been corrected here.

    Were Neandertals really extinct by 39,000 years ago?

    Now, in one sense, the survival of Neandertals after 40,000 years ago is not terribly important. Africans mixed with Neandertals, and as far as we can tell (an issue my lab is addressing now) the mixture is not preferentially within Europe. That argues for a West Asian interaction of the population, and it remains to understand why the ancestors of Europeans did not interact more than other populations. Probably a good hypotheses is that today's Europeans derive most of their ancestry from outside Europe during the last 10,000 years. If the Neandertals did not persist within Europe long during the Upper Paleolithic, that provides another alternative.

    But to say that we doubt a particular kind of information about dates is not the same as saying that Neandertals did not exist after 39,000 years ago.

    Direct dates on Neandertal bones are far from the only evidence of their persistence in Europe. Dozens of sites are dated by radiocarbon on fauna or charcoal. These dates themselves may be subject to the same critique as applies to the human bone. But there are not 15 of them, there are many, many more.

    For example, Gravina and colleagues [5] list 24 AMS dates for Châtelperronian contexts that are 36,000 BP or less. Calibration of dates for this era adds more than 3000 years or more to the calendar years represented by a radiocarbon date, so these are dates likely less than 40,000 years. They may be contaminated by recent carbon (and indeed a few are outliers below 32,000 BP), but if so some of them are remarkably consistent.

    Martínez-Moreno and colleagues [6] give a recent review of the Middle-Upper Paleolithic transition in Iberia. They list several sites with late Mousterian industries later than 34,000 radiocarbon years BP, even without counting the contentious examples (like Gorham's Cave) that arguably are later than 30,000 BP.

    I would not be happy assuming that every Mousterian site is a Neandertal site, not even in this limited geographic context. There is too much technical overlap, and sometimes small samples of artifacts, to be definitive about such an association. Technology is not biology. Neither would I be willing to assume that late Neandertals are entirely Neandertal -- we see no genetic evidence of African mixture into this population in the Vindija or El Sidron genomes, but these are older than 45,000 years. Who knows what a 35,000-year-old Neandertal in France or Spain (or Croatia) would look like genetically? But only Neandertal remains have thus far been associated with Mousterian and Châtelperronian in France and Iberia. Several sites have stratified Middle to Upper Paleolithic transitions with dates after 40,000 calibrated years BP.

    So from the Neandertal point of view, I think this is largely a non-story. There remains substantial question about the pattern of appearance of the post-Neandertal population, as I've extensively discussed here. When we consider the Caucasus, we are still working to understand the timing and mode of the later Neandertals and early Upper Paleolithic people. But there's really no serious challenge to the idea that Neandertals existed in Western Europe after 40,000 years ago.

    Or if there is, it'll be out of sync with what most of us think we know.


    References

    1. Hawks J, and Wolpoff MH. 2001. Paleoanthropology and the Population Genetics of Ancient Genes. American Journal of Physical Anthropology 114:269–272.
    2. Pinhasi R, Higham TFG, Golovanova LV, and Doronichev VB. 2011. Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus. Proceedings of the National Academy of Sciences [Internet] 108:8611–8616. Available from: http://dx.doi.org/10.1073/pnas.1018938108
    3. Higham T, Ramsey CB, Karavanić I, Smith FH, and Trinkaus E. 2006. Revised Direct Radiocarbon Dating of the {Vindija} {G1} {Upper Paleolithic} {Neandertals}. Proceedings of the National Academy of Sciences, U. S. A. 103:553–557.
    4. Blockley SPE, Bronk Ramsey C, and Higham TFG. 2008. The Middle to Upper Paleolithic transition: dating, stratigraphy, and isochronous markers. Journal of Human Evolution [Internet] 55:764–771. Available from: http://dx.doi.org/10.1016/j.jhevol.2008.08.009
    5. Gravina B, Mellars P, and Ramsey CB. 2005. Radiocarbon Dating of Interstratified {Neanderthal} and Early Modern Human Occupations at the {Chatelperronian} Type Site. Nature [Internet] 438:51–56. Available from: http://dx.doi.org/10.1038/nature04006
    6. Mart\'ınez-Moreno J, Mora R, and de la Torre I. 2010. The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula. Journal of Human Evolution [Internet] 58:211–226. Available from: http://dx.doi.org/10.1016/j.jhevol.2009.09.002
    Synopsis: 
    I disagree with a new story that claims Neandertals disappeared before 40,000 years ago.
  • Neandertal stories on parade

    Sat, 2010-12-04 23:21 -- John Hawks

    Long-time science journalist Robin McKie has a long article in The Observer about the Neandertals this weekend: "Neanderthals: how needles and skins gave us the edge on our kissing cousins".

    The article puts together several aspects of recent inquiry into Neandertal biology -- the genome sequencing, the dating questions over Châtelperronian artifacts from Grotte du Renne, and some of Steve Churchill's work on projectile versus thrusting weapons. There's a real interesting mix of stuff here, some that I agree with and basically find uncontroversial, and other stuff that I find to be outlandish or unsupported by any evidence.

    For example, McKie talked to Brian Fagan, who has a new book out (Cro-Magnon) that tries to describe the human "edge" over Neandertals. A good topic, but this paragraph is completely misleading:

    But which specific traits gave us such an advantage that we were propelled to global glory at the expense of the Neanderthals? In the suite of behaviours that we evolved in Africa 150,000 years ago, what were the characteristics that really made a difference and can therefore be considered as defining human attributes? There are many candidates – complex language and superior memory, for example. However, among many scientists there appears to be consensus that imagination and opportunism were critical attributes.

    There is no "suite of behaviours that we evolved in Africa 150,000 years ago." There just aren't any. There's no good evidence of symbolic expression, no projectile points, no subsistence innovations, no evidence of long-distance raw material procurement or trade. That's the big problem we have substantiating a modern human advantage -- the "modern" humans didn't seem to get many behavioral innovations in Africa that the Neandertals didn't get, and the Neandertals got them almost as early.

    It is an undeniable problem; there's no sense glossing over it. Churchill's (and John Shea's) ideas about projectile weapons are right now among the most reasonable suggestions, because there do seem to be relatively early (ca. 85,000-90,000 year old) projectile points in Africa.

    It would be convenient if there were better evidence that projectiles were a singular innovation. But as John Shea [1] wrote in 2006, the idea of projectile weapons seems to have gotten around widely, possibly including Neandertals:

    The evidence currently available instead favors an indigenous origin for projectile point technology in the Levant ca. 40–50 Ka. Similarly, the earliest European Upper Paleolithic stone artifacts that fit the TCSA criteria for projectile points, Chatelperronian points, Font Robert points (as well as Aurignacian split-based bone/antler points) do not have clear chronological antecedants in the Levant (though it is possible that other as-yet-unidentified projectile point types do). While it is possible that over-production of atmospheric radiocarbon between 30 and 50 Ka [39] obscures rapid geographical diffusion of projectile point technology the typological variability of the earliest likely stone and bone projectile points in Africa, the Levant, and Europe do not currently support a diffusion/migration hypothesis. It is vastly more likely that projectile point technology was developed convergently among African, Levantine and European hominin populations.

    I probably wouldn't stretch so far as to say that the Châtelperronian Neandertals were using projectile weapons, even if the points are consistent with that hypothesis. But considering that a big element of McKie's story is the dispute over the Châtelperronian evidence of ornamentation (at Grotte du Renne), I think it's fair to remind people that those late Neandertals had a lot of things going on. All the skeletal associations with the industry are Neandertal, and there are multiple sites representing the interesting material culture elements.

    I've actually been stunned lately by the number of people who have asked me about the Grotte du Renne paper and it's "demolishment" of the case for Neandertal ornamentation. I say stunned, because people seem completely unaware of the substantial Mousterian record of pigment processing and use.

    My candidate for the most subtly controversial element of McKie's story: the opening passage about the Swanscombe skull:

    Many treasures [at the Natural History Museum] compete for attention, but there is one sample, kept in a small plywood box, that deserves especial interest: the Swanscombe skull. Found near Gravesend last century, it is made up of three pieces of the brain case of a 400,000-year-old female and is one of only half-a-dozen bits of skeleton that can be traced to men and women who lived in Britain before the end of the last ice age. Human remains do not get more precious than this.

    However, the Swanscombe find is important for another, crucial reason: the skull is that of a Neanderthal

    I say that's controversial because it asserts that this 400,000-year-old skull is a Neandertal. The case for Swanscombe as a member of the Neandertal lineage has been mostly chronological, not because it has any pattern of derived Neandertal morphology. There were people in Europe before the Neandertals, they had a subset of Neandertal features, and so they were plausibly early members of a Neandertal lineage. But the genetic work this year, discussed later in the article, argued that humans and Neandertals shared a common ancestral population only 250,000-400,000 years ago. If that's true, the chronology is all wrong for Swanscombe to be a Neandertal itself. Indeed, this chronology would not permit Swanscombe to be a member of a population exclusively ancestral to Neandertals.

    But what, then, is it?

    I think the chronology is wrong, and I doubt whether the evidence will soon let us distinguish gene flow from isolation at this time depth. There's not much sense talking about the "human-Neandertal ancestral population" when some Neandertals were ancestors of some humans.

    Still, the Middle Pleistocene European population focuses the problem. If Neandertals themselves had derived much of their gene pool from Africa in the Middle Pleistocene, as the genetic work has suggested, what does that mean for specimens like Swanscombe? And if we substantially lengthen the chronology of human diversification, what does that mean for Middle Pleistocene Africans?


    References

  • Les Rois revisited, and dental classification of other Aurignacian individuals

    Tue, 2009-06-02 10:43 -- John Hawks

    I pointed a couple of weeks ago to the Les Rois Neandertal paper by Ramirez Rozzi and colleagues.

    In the new article section of Journal of Human Evolution, Shara Bailey, Tim Weaver and Jean-Jacques Hublin have a paper that examines the Les Rois sample (among many others) in terms of dental discrete traits. Basically, they set up a discriminant function that can tell Mousterian Neandertals from later Upper Paleolithic people with around 89 percent classification accuracy, and then they applied it to Aurignacian and Châtelperronian dental remains. Here's what they concluded about Les Rois (p. 13):

    The analysis of the entire Les Rois sample (n = 15) showed that they have an overwhelmingly ‘modern’ signal. Fourteen of the fifteen individuals had high posterior probabilities of belonging to the Upper Paleolithic modern human group (>80%). This is not unexpected considering the teeth are associated with an Aurignacian industry (Dujardin, 2000) and most come from unit B, which has been dated to 28,715 ± 145 BP using AMS C14. One individual (Mandible B), however, was classified as Neandertal with a low posterior probability (54%) based on 13 traits.

    Ramirez Rozzi (pers. comm.) has recently suggested that Mandible B represents a Neandertal, based largely on the asym-
    metrical P4 together with some aspects of the corpus. Trinkaus (2007) has also argued that the specimens from Les Rois are mixed in morphology. While the P4 is asymmetrical, a large sagittal crack in the crown exaggerates this feature and the remaining aspects of the tooth are distinctively not Neandertal-like (it lacks a transverse crest and multiple lingual cusps: Fig. 6). In the end, we do not consider the posterior probability of 54% to be compelling enough to conclude, based on dental traits, that there were Neandertals present at Les Rois.

    Ramirez Rozzi et al. (2009) made their conclusions about the Les Rois specimens mainly based upon perikymata packing patterns, secondarily supported by tooth sizes and the dental nonmetrics used here. So they are really concluding the same thing about mandible B. What's interesting is that Ramirez Rozzi and colleagues find several other dental individuals from the earlier unit with similar enamel formation patterns, and which they claim are also Neandertals.

    In some senses, Les Rois is a good case study for how more complete specimens come to dominate the discussion to the exclusion of other fragmentary remains. Mandible B is itself a tiny piece of a skeleton, but because it has several different anatomical elements on that little piece, the two papers can conduct this kind of in-depth analysis. But the status of the other isolated teeth are equally important -- if Ramirez Rozzi and colleagues are right, they might be enough to establish the earlier Les Rois sample as standing apart from other early Aurignacian-associated samples. If we're talking about a single specimen, however well documented, the situation is somewhat different.

    As everybody knows, my null hypothesis is that the samples are mixed in their morphological pattern. On that topic, the more interesting implication of the study by Bailey and colleagues (2009) is that when they applied their discriminant function to Châtelperronian samples, they got a similar classification frequency to Neandertals as for earlier Neandertal samples. And when they applied their function to Aurigacian samples, they got a similar classification frequency to modern humans as for later Upper Paleolithic samples. Under the hypothesis of a mixed transition from earlier Neandertals through later Châtelperronian Neandertals into early Aurignacian and into the Upper Paleolithic, you'd expect the intermediate time steps to give a lower classification frequency -- more Neandertal-like in Aurignacian than later; more modern-like in Châtelperronian than earlier. Bailey and colleagues found that their function classified 85 percent of Aurignacian dental individuals as modern (29/35) compared to 89 percent of later Upper Paleolithic individuals (56/63). Those numbers aren't significantly different, and given the variability in completeness of assigned specimens, I wouldn't go farther.

    One possible criticism of the paper is that the morphological pattern that makes up the discriminant mostly consists of traits that are shared by both groups but differ in frequencies. It is still quite possible to use the traits to discriminate individual specimens, but somewhat harder to interpret what a change in trait frequencies means in genetic terms. Bailey and colleagues recognize this issue, and raise it in their discussion of the Oase 2 specimen, for example (p. 12):

    Given that the cranium of Oase 2 is clearly not that of a Neandertal (Rougier et al., 2007), the assignment of this individual to the Neandertal group was unexpected. Trinkaus (2007) has suggested that, while essentially ‘modern,’ both Oase 1 and 2 exhibit a mosaic of cranio-dental features, some of which are archaic (e.g., dental proportions, long and flat frontal bone), and others apparently derived towards anatomically modern humans (parietal curvature, absence of supraorbital torus) or towards Neandertals (unilateral lingual bridging of the mandibular canal).

    It is important to note that the dental traits aligning Oase 2 with Neandertals are archaic in nature, as they are observed in other fossil hominins as well (Bailey, 2002b, 2006; Martínon-Torres et al., 2007). It is unfortunate that incisor morphology could not be assessed (teeth are missing), and that the upper M1s are too worn to ascertain occlusal polygon shape and occlusal polygon area, since these are features that are likely derived for the Neandertals/Neandertal lineage (Bailey, 2004; Gómez-Robles et al., 2007). Considering that some of the most diagnostic features of the upper dentition could not be assessed and that our approach is not 100% accurate, we caution against over-interpreting the classification of Oase 2.

    That's the basic problem of comparing Neandertals with humans, which we encounter genetically as well as morphologically. The groups differ in the frequencies of traits, but not often in the exclusive presence of distinctive ones. Sometimes, what once looked like a distinctive trait is then found in the other group -- so it's not distinctive anymore! Some researchers focus on "distinctive combinations" of traits, which tend to include a mixture of primitive and derived morphologies. But differences in trait frequencies automatically lead to differences in the combination of traits. Sometimes combinations are disproportionately represented (that is, putting traits together separates the samples more than considering them individually), but it is unclear how much of a trait combination may be explained by a history of inbreeding (in isolated populations) and how much may be explained by pleiotropy (of a few genes that differ in frequency).

    You'd think this problem would be easier than it is. But look at Les Rois -- a fragmentary but interesting sample, with a blend of morphologies in different specimens. How do we interpret the similarity of Les Rois mandible B to Neandertals? Is it a Neandertal? If we considered the dental nonmetric features alone, as Bailey and colleagues suggest, the specimen looks like a Neandertal but with really rather weak evidence. If we add the perikymata and size data, the similarities with Neandertals are increased, and other teeth from the site also tend to look more Neandertal-like. But we know that perikymata patterns vary in recent human populations. How should we consider this variation as we compare the Les Rois teeth -- should we consider Europeans only, or modern humans more broadly? How did those traits change within the last 30,000 years, and is that relevant to the 10,000 years before?

    Well, it's certainly enough to keep things interesting. I'm raising a lot more questions than offering answers. I like the approach Bailey and colleagues have taken because it includes much of the available sample in a way that can be considered as a unit. But then when we return the the issue of particular specimens and the possible patterns of genetic causation of the traits, we are left with the same problems as before.

    References:

    Bailey SE, Weaver TD, Hublin J-J. 2009. Who made the Aurignacian and other early Upper Paleolithic industries. J Hum Evol (in press) doi:10.1016/j.jhevol.2009.02.003

  • "Competitive exclusion" and the extinction of Neandertals: should we believe it?

    Tue, 2008-12-30 23:44 -- John Hawks

    I've been out of e-mail range for the past week. In the meantime several people e-mailed me this new paper:

    Neanderthal Extinction by Competitive Exclusion

    Background: Despite a long history of investigation, considerable debate revolves around whether Neanderthals became extinct because of climate change or competition with anatomically modern humans (AMH).

    Methodology/Principal Findings: We apply a new methodology integrating archaeological and chronological data with high-resolution paleoclimatic simulations to define eco-cultural niches associated with Neanderthal and AMH adaptive systems during alternating cold and mild phases of Marine Isotope Stage 3. Our results indicate that Neanderthals and AMH exploited similar niches, and may have continued to do so in the absence of contact.

    Conclusions/Significance: The southerly contraction of Neanderthal range in southwestern Europe during Greenland Interstadial 8 was not due to climate change or a change in adaptation, but rather concurrent AMH geographic expansion appears to have produced competition that led to Neanderthal extinction.

    OK, so should we believe it?

    The authors are out to test the idea that climate killed the Neandertals (also covered by me in 2007, not to mention "The unbearable hotness" from last week).

    The authors confine their analysis to a simple question: Were the European ecologies of the later times of Neandertal existence compatible with those that existed slightly earlier? If the climate deterioration is insufficient to explain the range reduction of late Neandertal sites, then we need some other factor. Competition with the non-Neandertal population would be a logical hypothesis, in this event.

    Competitive exclusion is not a new concept applied to Neandertals. The novel element in the paper is its inclusion of paleoclimate models to support the hypothesis that Neandertals and modern humans actually would have competed in the same niche.

    The authors applied an optimization algorithm to data. The data included:

    1. Paleoclimate predictions for small areal units within Europe for three time periods between 43,000 and 35,000 (calibrated) years ago.

    2. The locations and dates of archaeological sites dating to these periods, whether Mousterian, Châtelperronian, or Aurignacian. The first two are assumed Neandertal, the third modern human.

    The algorithm tries to find shared paleoclimate features among the locations represented by archaeological sites. Once these are found, the algorithm finds other areas that fit the same paleoclimate parameters as those where archaeological sites were found. In other words, it is an attempt to determine the total ecological range of the populations represented by the sites.

    For example, here are the results for the earliest of the three time periods, H4. This is a comparison of the maps of Europe for both the Mousterian-Châtelperronian (left) and Aurignacian (right) archaeological samples:

    H4 paleoclimate predictions for Neandertals and modern humans

    In this map, red areas are those predicted to be suitable for habitation by Mousterian-Châtelperronian (left) and Aurigacian (right) populations, respectively. One thing stands out: they have almost identical ecological tolerances. Modern humans were not using different ecological zones than Neandertals. The analysis of the later periods shows that the modern humans were not exploiting climate changes at the expense of Neandertals.

    The key graph of the paper shows that during the latest time period, near 35,000 years ago, the paleoclimate models predict a very large area of Europe would have been suitable for Neandertal habitation -- at least, if their habitation were constrained only by climate. But the Neandertal sites during that time period are restricted to a very small area. So some additional factor is required. The authors promote the hypothesis that the important factor was the population growth of modern humans.

    Now, should we doubt the results? I think this is a good test of the hypothesis that climate change killed the Neandertals. It didn't. They survived through an entire glacial cycle before 40,000 years ago. Without some other factor, they would still be here today.

    The paper is stronger than many that have tried to make a similar argument -- that climate couldn't have killed various extinct megafauna. In large part, that is because both the American megafaunal disappearances and the entry and growth of human populations coincided with a period of rapid climate change. In the time frame of the last Neandertals, there were important climate changes, but the paleoclimate models indicate that these changes weren't enough to make Europe uninhabitable for either humans or Neandertals.

    The paper is not a test of Neandertal genetic extinction. It takes Neandertal population disappearance as a given. Models that involve gene flow or cultural exchanges between Neandertals and other populations are not part of this paper's scheme.

    In this sense, the assumption that the archaeological industries can be analyzed with methods developed for species seems questionable. The paper acknowledges this issue:

    Our assumption is that human adaptive systems, defined here as the range of technological and settlement systems shared and transmitted by a culturally cohesive population within a specific paleoenvironmental framework, can be considered to operate as a ‘species’ with respect to their interaction with the environment. This does not imply, however, that human adaptive systems necessarily remained stable over time, as might be the case with animal species occupying narrow and stable niches. Humans can change their adaptive systems rapidly through technical and social innovations in response to environmental change. We know, however, that this was not the case during the late Middle and Upper Paleolithic, periods during which specific human adaptive systems spanned a number of climatic events. Thus, the method described in this study is particularly relevant for addressing issues of human adaptive system stability and eco-cultural niche stability (Banks et al. 2008:2).

    But Neandertals during the span from 43,000 to 35,000 years ago were adopting various Upper Paleolithic technological elements. That seems to contradict the assumption that the "technological and settlement systems...operate as a 'species'." Instead, it seems to indicate that these systems changed significantly across the time frame modeled in the paper. I don't think that observation weakens the hypothesis that modern and Neandertal populations may have competed in the same ecological niche. If the Neandertals were using the same technical elements, it probably reinforces the hypothesis of competition.

    But I think it is important to bear in mind what the paper tested. The analysis rejects the hypothesis that climate was sufficient to drive a range restriction of Mousterian and Châtelperronian. It doesn't provide additional information about the mode of such restrictions. With that in mind, I admire the paper and see some useful additional work that might be tackled with similar methods.

    References:

    Banks WE, d’Errico F, Peterson AT, Kageyama M, Sima A, et al. (2008) Neanderthal Extinction by Competitive Exclusion. PLoS ONE 3(12): e3972. doi:10.1371/journal.pone.0003972

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Neandertals

For years, I've worked on their bones. Now I'm working on their genes. Read more about the science studying these ancient people.

Denisova

From a finger bone of an ancient human came the record of a completely unexpected population. My lab is working on the science of the Denisova genome.

Acceleration

The advent of agriculture caused natural selection to speed up greatly in humans. We're uncovering some of the ways that populations have rapidly changed during the last 10,000 years.

Malapa

Just outside Johannesburg, the Malapa site is producing some of the most exciting finds in human evolution. This site is the headquarters of the Malapa Soft Tissue Project.