john hawks weblog

paleoanthropology, genetics and evolution

Homo erectus

  • Martin versus Falk on microcephaly

    Thu, 2006-05-18 23:45 -- John Hawks

    Science is carrying an exchange of technical comments about microcephaly and the endocast of LB1. Bob Martin and colleagues weigh in with an argument for why LB1 was a microcephalic. Dean Falk and colleagues respond that the microcephaly explanation can't account for the endocast's features.

    So what's the real story?

    Why isn't it obvious that this specimen was microcephalic?

    To some, it is a tautology: a microcephalic has a brain size significantly less than the normal range; this skull had a significantly small brain (near the minimum for Australopithecus!); ergo, this was a microcephalic.

    But microcephaly is a rare condition, and it is not a single condition, but a spectrum of conditions. There is a long chain of developmental processes leading to the normal human brain, including genes and necessary nutritional and chemical conditions. A break or interruption in many of these may result in significantly smaller than normal brain size. Many of these cause different anatomical configurations -- brains of different shapes. The endocast of LB1 was broadly normal in shape (at least, compared to early Homo) but very small in size.

    What seemed essential was to find a sample of modern human microcephalics that had the shape of LB1. It would be even better to find additional cranial or postcranial features of microcephaly that match the skeleton of LB1.

    An earlier comment by Weber and colleagues (discussed in this post) presented a comparison with a sample of microcephalics, which easily matched the LB1 endocast in size, but not in shape. The closest endocasts in shape still presented some differences with LB1 that stand out as divergent compared to normal modern human endocasts and early hominid endocasts.

    It's sort of like looking for a match with a tan Toyota Corolla, and they found a tan Honda Civic -- close in shape, but some telling differences.

    Now, at this point it seems like an exercise in pattern matching. Folks are going to look through museums at microcephalics until they find one that's closer to LB1.

    Did Martin and colleagues find a better microcephalic?

    Here's what Martin et al. write:

    Microcephalic skulls and endocasts similar to LB1 include the specimens shown in Fig. 2. Doubling of the volume for half-skull B yields a cranial capacity of 432 cc, close to that of LB1. Specimen C has a volume of 340 cc. Both lack obvious pathologies. For example, the cerebellum is tucked under the cerebrum (3).

    That gets around one of the criticisms of the close endocast presented by Weber and colleagues (2005). Is it enough?

    Falk and colleagues reply that the line drawings provided don't give enough detail to tell whether the "matching" endocasts actually are comparable with LB1.

    There are two ways to look at this. Here's an analogy: we've found a tan Toyota Corolla, but it isn't a tan Toyota Corolla with a dent in the trunk.

    The importance of the similarity depends on the question we want to answer. If we want to know if the cars were made by the same manufacturer, then we don't care about the dent, or even the color.

    If we want to know if our key will fit, then it matters a lot -- and we may need to know even more, like the license plate number.

    Falk and colleagues are looking for the license plate number:

    Because of brain shrinkage, one would also not expect to obtain a highly convoluted endocast (like LB1's) from such a specimen (5, 6), and we gather from the lack of detail on Martin et al.'s line drawings that neither of their microcephalics reproduced endocasts that are highly convoluted.

    ...

    Although the authors provide a line drawing for LB1's skull, no image is provided for its endocast. A line drawing is provided for an endocast of a microcephalic from the Field Museum, but not for its skull, which, as described (and illustrated in actual photographs) in the reference cited by Martin et al., "is long-headed and narrow, with a lowly vault, the face narrow, with apelike protrusion of the jaws" (8). This skull differs starkly from LB1's, which is extremely brachycephalic (2, 9).

    It would be helpful if we knew in advance which features were important ones for the comparison. The size is not enough, but is the shape? Or the shape and the convolutions? Or Brodmann's area 10? It seems a bit much to suppose that a skull has to share every feature of LB1 to count as the same developmental problem. We don't demand that of any other kind of diagnosis in living people. But we do demand that a key set of symptoms be present for a diagnosis, and in this case we don't have any guidance about which are the important features.

    Yuck.

    What is all this about Jakob Moegele?

    Martin and colleagues went into museum records to figure out the identity of the microcephalic included in the study by Falk et al. (2005a). This research showed that endocast was probably not very comparable to LB1 -- the skull belonged to a 10-year-old boy, it was a bad cast, it was a very small endocast even compared to LB1, etc.

    Falk and colleagues reply that it might not matter so much that the boy was only 10 because the brain size is essentially complete, older than the median microcephalic death age, not unusual in shape compared to other microcephalics, etc.

    I think there is little light here. Clearly no answers are possible from a single microcephalic. This microcephalic was not like LB1, but there are many microcephalics -- probably most -- that were not like LB1.

    Indeed, I wouldn't rule out that LB1 was microcephalic even if no modern microcephalics have its exact characteristics, since no modern microcephalics lived relatively long lives in Stone Age societies. There is a component of variability in examining such ancient skeletons that cannot be corrected with comparisons of living and recent humans. This may be a part of that.

    How about the dwarfing allometry stuff?

    Martin and colleagues present an argument for why the brain size of LB1 could not result from dwarfing:

    The tiny cranial capacity of LB1 cannot be attributed to intraspecific dwarfism in H. erectus. Body size reduction in mammals is usually associated with only moderate brain size reduction. Starting from three potential ancestral forms (H. erectus broadly defined; the chronologically and geographically closest H. erectus specimens from Ngandong, Java; and the substantially earlier Dmanisi hominids from Georgia) and following a range of possible dwarfing models, the predicted body size of a dwarf hominid with the cranial capacity of LB1 ranges from less than 1 g to 11.8 kg (Table 1 and Fig. 1) (4). Most of the figures calculated are at least an order of magnitude smaller than the estimates for LB1 (16 to 29 kg) (1). The largest are based on the insular dwarfing of elephants on Mediterranean islands (Model A) from 10,000 to 15,000 kg down to 100 kg. Despite the extreme dwarfing involved, and the relatively steep brain-body size scaling slope, the predicted body size for the dwarf hominid is still unrealistically small. Typical mammalian intraspecific scaling (Model B) indicates a maximum body weight less than half that estimated for LB1. Intraspecific brain-body size scaling in primates, including humans, is notably flat, particularly for males and females separately (5). This model (Model C) predicts tiny body weights for LB1.

    There are also an associated figure and table. This critique is correct, and close to close to what I wrote in my 2004 FAQ on the specimen. You can't accuse me of leaving out the significant details!

    But Falk and colleagues aren't hemmed in by this critique, because they aren't committed to the position that LB1 was a phyletic dwarf. For all they know, it could be an australopithecine -- in which case, the brain would not be unusual. That hypothesis presents its own problems (how did they get to that island, again?) but the brain size is certainly not one of them.

    What do I think?

    For the most part, my opinion from the October 2005 update still holds. I still think the specimen has widespread pathology, and I still think that's sufficient to question whether it was representative of its population.

    But... I have a new hypothesis I've been working on. It's secret. OK, so now I am leaving out significant details....

    Personally, I can't wait to see the long critical papers come out. Then we'll have a better idea of the boundary conditions.

    I think that Carl Zimmer makes a great point in his post this week:

    I wondered [in October], and I wonder now, why the editors of Science don't make sure that everyone agrees on the ground rules for comparing these brains before they publish? Otherwise both sides just squabble about methods and presentation, rather than about meatier matters.

    The problem may be that in both cases Science has relegated this exchange to the "Technical Comment" section, where reports are much shorter than normal papers. The descriptions of methods used in the research are often scant, and the comments also tend to include cryptic interpretations that cry out for more explanation. Falk and her colleagues say that the Hobbit's brain is consistent with apes or australopithecines, not Homo erectus. Now, I'd imagine that this might imply that the Hobbit descends not from Homo erectus, but from some Australopithecine that came out of Africa. That would be huge news if true. Yet the scientists just leave us hanging with a statement that is so cryptic as to be nearly useless.

    They must face this problem a lot, though. Who is Science going to bring in as a referee to make the two sides agree? And are they willing to let it go to another journal? Because these folks will take their work elsewhere rather than be forced to use someone else's methods.

    Personally, I think they should have a special hobbit issue where they hash out all these things. If they can have special Drosophila genome issues and special Cassini issues, they can surely have a special hobbit issue.

    I'd be happy to contribute a paper!

    References:

    Falk D, Hildebolt C, Smith K, Morwood MJ, Sutikna T, Jatmiko, Saptomo EW, Brunsden B, Prior F. 2006. Response to comment on "The brain of LB1, Homo floresiensis". Science 312:999. DOI link

    Falk D et al. 2005a. The brain of LB1, Homo floresiensis. Science 308:242-245. Full text (free)

    Falk D et al. 2005b. Response to comment on "The brain of LB1, Homo floresiensis". Science 310:236. Full text (subscription)

    Martin RD, MacLarnon AM, Phillips JL, Dussubieux L, Williams PR, Dobyns WB. 2006. Comment on "The brain of LB1, Homo floresiensis". Science 312:999. DOI link

    Weber J, Czarnetski A, Pusch CM. 2005. Comment on "The brain of LB1, Homo floresiensis". Science 310:236. Full text (subscription)

  • Hobbit news from Stony Brook

    Thu, 2006-05-18 23:45 -- John Hawks

    In another post I write about the Martin-Falk exchange on the microcephaly issue.

    Here, I review the Paleoanthropology Meetings summary by Elizabeth Culotta in Science.

    Along with some background, the article basically covers two talks: one by Susan Larson concerning the humerus, and one by Bill Jungers about the pelvis.

    Larson concluded that the upper arm and shoulder were oriented slightly differently in H. floresiensis than in living people. The shoulder blade was shrugged slightly forward, changing its articulation with the humerus and allowing the small humans to bend their elbows and work with their hands as we do. This slightly hunched posture would not have hampered the little people, except when it came to making long overhand throws: They would have been bad baseball pitchers, says Larson.

    When Larson looked at other human fossils for comparison, she found another surprise: The only H. erectus skeleton known, the 1.55-million-year-old "Nariokotome boy" from Kenya, also has a relatively untwisted humerus, a feature not previously noted. Larson concluded that the evolution of the modern shoulder was a two-stage process and that H. erectus and H. floresiensis preserved the first step.

    Humeral torsion was the feature that previously led to the (wrong) idea that the hobbits were quadrupeds, but that wrong idea just shows how odd the feature is in the context of hominids.

    In that context, you might wonder how it could be missed in a well-known specimen like KNM-WT 15000. Well, the picture helps explain:

    LB1 (top), human (middle), KNM-WT 15000 (bottom) humeri. Sized to approximately equal lengths, not to scale.

    Hmm.... I know I wouldn't want to be in that position -- it's a tough assessment to make on that specimen. The head might have been slightly posteriorly oriented there, but the torsion in LB1 is pretty low (low being strange in this comparison) -- so I would be really hesitant to match them up. The problem is that identifying the feature really takes a fairly complete humerus, and there aren't very many either from Australopithecus or early Homo. As reconstructed, AL 288-1 (Lucy) doesn't have low torsion, and I wouldn't have assessed Sts 7 as having it either, although there is some possible distortion there. So it wasn't an ancestral feature in hominids, so far as we can tell. Without an epiphysis, KNM-WT 15000 lacks some of the most relevant anatomy, but there might be enough....

    I'll really look forward to seeing more detail about this argument -- it should be interesting to see the anatomical comparisons!

    The other talk was Bill Jungers', which focused on the pelvis:

    In a separate talk, Jungers reported more unexpected findings. He was able to reconstruct the pelvis, which had been broken when the bones were moved to a competing lab in Indonesia (Science, 25 March 2005, p. 1848). Although previous publications had described the pelvis as similar to those of the much more primitive australopithecines, Jungers found that the orientation of the pelvic blades is modern. The observation adds weight to the notion that hobbits had H. erectus, rather than australopithecine, ancestry.

    This should be interesting too -- there has been a lot of disagreement about how flat australopithecine ilia actually were. Many of the most well-known fossils are very flattened, but there is also a lot of distortion. Hopefully any distortion introduced by the breakage (pictures in earlier post) hasn't affected these comparisons.

    The spin on these stories is that they make it more likely that the Flores hominids are descendants of early Homo rather than Australopithecus. Personally, I would guess that the extent of dwarfing necessary to explain the size of the specimen pretty much throws its postcranial affinities up for grabs. For example, it's never been clear how many of the Australopithecus-Homo anatomical differences are merely consequences of body size, and how many reflect different adaptations. Since so many of the apparent differences are at least responsive to allometric constraints, it's a problem that always has to be faced.

    Let's put it this way: if it weren't a problem distinguishing these genera from postcrania, there would be a lot less confusion assigning the unassociated Koobi Fora postcrania. For LB1, it helps that the bones are complete -- but, hey, we can't even tell yet that it's not a pathological modern human!

  • Hoffmeyer on language as adaptation

    Fri, 2006-05-12 16:04 -- John Hawks

    Chapter 8 of Hoffmeyer's Signs of Meaning in the Universe is about the evolution of language. I really like the opening paragraph, which is worth remembering:

    I have observed that many of my students -- who are, of course, studying to become biologists -- are extremely reluctant to accept the idea of human language as something special. They point out that animals such as dogs, whales or chimpanzees might well have a language that we human beings have just been too highfalutin to acknowledge. And, in my experience, my holding up of the novels of Dostoyevsky or the Bible as examples of how human language is something quite unique in this world seems to make no great impact. If anything, I have the feeling that these students look upon humanity, and human nature in general, as a warped work of nature epitomized by its destructive penchant for building concrete blocks of slums and waging war. And, seen in that light, the fact that a few sensitive individuals might find it in themselves to write emotionally harrowing works of fiction does not seem all that strange.

    I have to say, I have had the identical experience with students -- down to their arguing that animals may have their own versions of Shakespeare. "Who is to say they don't?" is the argument I have heard a lot.

    My reaction is different than Hoffmeyer's, though. For one thing, these students have a long upbringing of being told it is wrong to make value judgments; for another, they have a long upbringing of watching movies with talking animal characters who emote with real human feeling.

    I surmise it is difficult for some to imagine how any animal could exist without human-like mental capabilities.

    As a corollary, I often ask my classes whether they would rather be hunted by a human or by a chimpanzee. I have never yet had a class where more than a few students choose the chimpanzee -- even though a chimpanzee would be enormously less dangerous as an enemy than a human. (For one thing, a chimpanzee is rather less likely to continue to track you after dark...) It seems to come down to a fear of the unfamiliar, and the optimistic idea that you could reason with a human enemy.

    I choose not to worry until the army breeds a legion of warrior chimps.

    In any event, Hoffmeyer cites Merlin Donald (1991) for much of his scenario for language evolution. The basic idea was that there was an intermediate stage between apelike minds and human language that was based on mimetic abilities, "the ability to carry out collective motor-based reconstructions of earlier incidents." Mimetic culture required and built upon social intelligence (and associated social learning abilities), but it also required a link between episodic memory (remembering events) and procedural memory (remembering how to carry out an action). Ultimately, these led to an internal syntactic memory that helped organize conscious thought, a system from which language sprung.

    Under this scenario, mimetic culture characterized Homo erectus, who lacked language, but shared most other mental characteristics with humans.

    This passage raises an interesting idea:

    It is a fascinating thought that Homo erectus, with a brain capacity not really so far removed from the present dimensions or our own brains, may have been very much like us in almost all respects -- and, especially from an emotional point of view, in the most profound ways. Why, were it not for that one little quirk -- the lexicon, that ability to send our inner experiences flowing from our lips in streams of words to be pondered and debated among ourselves, adopted or rejected -- we might be said to have been almost identical at birth. The difference between the talking Homo sapiens and Homo erectus may not have been any greater than that between the people of the later Stone Age and the people of today. Because what separates modern man from Stone Age man is the existence of external (extrasomatic) memory banks -- first and foremost the written word in the form of books, but also the legacy of sculptures, pictures, buildings, tools and, these days, computers. The presence of these external memory banks implies that we, as adults, bear the burden not only of our own inherent intellectual legacy but also of a hundred-generation-long struggle to extract the essence of our forefathers' experience. This struggle has taught us to live in a world saturated by science, technology and art -- a world which could quite conceivably create an even greater gap between us and the mind of Stone Age man than Stone Age man, by virtue of the spoken word, created between himself and the mind of Homo erectus.

    Now, this seems vital to me: is the extent of human variation today -- a result of recent genetic and cultural differentiation -- as great as the difference between Homo erectus and modern humans? Hoffmeyer considers the problem in terms of culture alone, and concludes that the difference might well be as great.

    In support of this, he suggests that the early effects of culture on brain development generate possibly vast phenotypic differences among humans:

    [Differences in programmed cell death] results in the cultural stamp leaving a telling imprint on the neurological structure of the brain. If not at birth then certainly by the time they start school modern human beings are therefore already very different from the people of the Stone Age. Just to be on the safe side it ought also to be mentioned that this restructuring of the neurological terrain is not altogether irrevocable. It appears at any rate that, even in adults, the area of the cerebral cortex which registers hand movements can be expanded or reduced as required. But for anyone desirous of reverting to the Stone Age mentality it would hardly be enough just to journey back to the settlement at Vendsyssel-Thy. You would have to retreat pretty far into the Siberian taiga and stay there for years, and in fact it would be best to start out in early childhood.

    Of course, there are modern humans living in such contexts today. What about them? The "Stone Age - modern" difference is not merely a temporal comparison (prehistoric vs. today), it is also a geographic and cultural comparison (civilization vs. hunter-gatherer). From this perspective, Hoffmeyer's point easily misfires -- the claim would seem to require that the integration of a Homo erectus-like human into a modern hunter-gatherer group should be no more difficult than the integration of today's hunter-gatherers into civilization.

    But modern hunter-gatherers do have many problems integrating into larger societies. The ability to speak is not particularly a barrier, but linguistic differences are. Culture, technology, and economics are all barriers. Probably the most severe barrier is disease.

    Archaeologists have a long history of using modern hunter-gatherers as analogues for ancient humans precisely because of their differences from larger societies in terms of economy, subsistence, and social organization. That analogy assumes that the gulf between Homo erectus or other ancient hominids and modern hunter-gatherers is not so great. This tradition has emphasized certain effects of language and symbolic culture, such as an increase in the possible size of social units, breadth of economic and trade relationships, and sophistication of technology.

    Modern and historic hunter-gatherers are diverse in these aspects of behavior -- everything from marriage patterns, kinship ties, and food acquisition strategies to land tenure and dispersal strategies. Certainly such diversity itself would be impossible in the absence of language: could different kinship systems be possible in the absence of the linguistic structures that support them? Or different taboos?

    Two perspectives present themselves. One point of view would see the variation among modern humans as highly significant; a clear indicator of symbolic culture and language and a strong element in shaping diversity in the minds of modern humans. The other point of view would see the minds of modern humans as essentially similar despite any cultural differences, in which case symbolic culture and language can have had little effect on minds outside the relatively narrow parts driving symbolic culture and language.

    The two perspectives are very different in their predictions about differences among modern human minds. But they are essentially the same with regard to the archaic-modern human differences compared to modern human variation.

    References:

    Donald M. 1991. Origin of the Modern Mind: Three Stages in the Evolution of Culture and Cognition. Harvard University Press, Cambridge MA. Amazon

    Hoffmeyer J. 1996. Signs of Meaning in the Universe. Indiana University Press, Bloomington IN. Amazon

  • "Spacecraft all over the Pliocene"

    Thu, 2006-04-27 08:34 -- John Hawks

    Rex Dalton has a great two-page article in Nature about the bush vs. ladder dispute. It keys off of the Middle Awash Australopithecus anamensis article by White and colleagues from a couple of weeks ago.

    If you recall that one, White et al. posited that Ardipithecus was likely ancestral to Au. anamensis, and that the two did not overlap in time. Here's the key exchange in the Dalton piece:

    This month's Nature paper makes a bold argument, and shows the Awash team seeking to put its mark on the record. Others in the
    field are impressed. "When you find 30 new hominid fossils, you are allowed a certain amount of conjecture," says Bernard Wood, a palaeoanthropologist at George Washington University in Washington DC. "As always, they have done a fantastic job."

    But he and others are unconvinced by the Awash team's conclusion: "This is only the first half of the rugby match," says Wood. Meave Leakey, lead author on the Au. anamensis discoveries in Kenya, is more blunt. "I don't believe this," she says. "We do not have the specimens to fill the gaps."

    Leakey and Wood are among those who believe that other, as yet undiscovered hominid species may have lived at this time, from 4.4 million to 2.9 million years ago. The existence of other species would cloud or eliminate the argument for a direct lineage. "My prejudice is there are more lineages rather than fewer -- more diversity," says Wood. "I have to concede these new data are dramatic. But we should beware coming out with a complete explanation when we don't have all the
    evidence."

    This argument frustrates White. "There were Martians there back then too," he says. "And spacecraft all over the Pliocene -- we just haven't found them yet."

    Waiting for Monte Cassino

    In a series of articles since 2000, White and colleagues have laid out a systematic attack on the "bushy" phylogeny model. Their arguments have extended across four million years and seven species, with a breadth that rivals the Allies breaking the Winter Line.

    Consider the angles of attack:

    1. Au. anamensis -- Au. afarensis. Everyone basically accepts that Au. anamensis is a direct ancestor of Au. afarensis. And the two species are really not very different from each other -- for instance, they are more alike than either is to Ardipithecus. The transition between these species would look to be a simple case of anagenesis, except...

    ...for Kenyanthropus (Leakey et al. 2001). This small-toothed, flat faced hominid needs an ancestor, too. Au. anamensis might have been the common ancestor of Kenyanthropus and Au. afarensis. If so, then both these later species originated by cladogenesis from Au. anamensis. A similar argument might be made for other species, like Australopithecus bahrelghazali (Brunet et al. 1996) or the Sterkfontein Member 2 hominids. But Au. bahrelghazali is only known from a partial mandible and only differs from Au. afarensis by a three-rooted premolar, which is considered by many to be weak evidence, and the Sterkfontein Member 2 sample has not yet been taxonomically assigned -- they might turn out to be Au. afarensis, for example. Kenyanthropus remains the strongest case for cladogenesis (i.e., a "bush"). Yet...

    ...White (2003) denied that the Lomekwi skull KNM-WT 40000 was a distinct species. In particular, he argued that the extensive postmortem deformation of the skull made it impossible to substantiate an anatomical difference from Au. afarensis, and even if it was different, the anatomical diversity of living hominoid species is so great that it would probably encompass the difference between KNM-WT 40000 and known Au. afarensis crania.

    2. Earliest hominids. At the moment, the earliest putative hominids include three genera: Orrorin (Senut et al. 2000), Sahelanthropus (Brunet et al. 2002), and Ardipithecus, represented in the Late Miocene by Ar. kadabba (Haile-Selassie 2001, Haile-Selassie et al. 2004). Evidence for obligate bipedality has been challenged (by different researchers) for each of these three (I'm one of those who has questioned bipedality for Sahelanthropus).

    So far the only comparable anatomical parts from all three samples are teeth...

    ...which were examined by Haile-Selassie, Suwa and White (2004). They concluded that the variation among these three genera

    is no greater in degree than that seen within extant ape genera. Despite claims of molar enamel thickness differences among these late Miocene fossils, we question the interpretation that these taxa represent three separate genera or even lineages. Given the limited data currently available, it is possible that all of these remains represent specific or subspecific variation within a single genus (Haile-Selassie et al. 2004:1505).

    Additionally, Ohman, Lovejoy and White (2005) challenged the interpretation of the internal anatomy of the Orrorin femur, which had been suggested to be more derived than that of Au. afarensis. They wrote:

    We agree that the Lukeino femur's external morphology suggests some form of bipedality. Yet the more detailed original scans appear to show a distinct superior cortex different from Australopithecus and humans, with the cortex distribution being more primitive than that seen in any other hominid, including Australopithecus.

    The relevance of this argument to the phylogenetic diversity of early hominids depends on the anatomy of the Ardipithecus femur, which none of the rest of us are in a position to know. But one may speculate that if all these early "hominids" had femora with similar morphology, it would further reinforce the interpretation that they belong to a single lineage.

    3. Ardipithecus -- Au. anamensis. This is the current example. Here's how Dalton discusses it:

    The latest Afar discovery is exciting experts because it shows that the three hominids existing in the same area, but in successive time periods. Tim White of the University of California, Berkeley, co-leader of the Awash team, believes this points to a direct lineage between the three -- a process called phyletic evolution. The new Au. anamensis fossils are only 300,000 years younger than Ar. ramidus, meaning that if one became the other, the changes would have had to happen that fast. But the key point, says White, is that fossils of Au. anamensis and Au. afarensis have never been found in sediments the same age as those containing Ar. ramidus. If fossils of the different species were found together, that could show that they belonged to multiple lineages existing simultaneously.

    Finding remains of all three species in the same area but not from the same time period suggests they did not coexist, says White.

    ...

    The specimens also provide anatomical clues to evolutionary history. "The new Au. anamensis fossils are anatomically intermediate between the earlier Ar. ramidus and the later Au. afarensis," says White. For example, the teeth of the newly discovered Au. anamensis fossils seem adapted to chew tougher and more abrasive foods than Ar. ramidus. The researchers believe this shows that Au. anamensis had a broader diet. "All this strengthens the view that there is phyletic evolution from Ar. ramidus through Au. anamensis," says White. He believes he has nailed down the relationship between the two later species, although he says that further specimens are needed to prove the earlier link (Dalton 2006:1100).

    Of course, it would help matters if we knew in more detail what Ardipithecus looked like. But one must imagine that the stage is being set for its revelation. The unilineal interpretation places Ardipithecus at the critical point as an ancestor to the major mid-Pliocene australopithecine lineage. Extending the unilineal interpretation earlier into the Late Miocene would make Ardipithecus the earliest hominid as well.

    It is not necessary to think that taxonomic uniformity means anatomical uniformity, though. Ardipithecus already encompasses a trend of decreasing canine size and less sectorial P3 for example. A trend toward fuller skeletal adaptation to bipedality may also be imagined. But in that context, it is important to note that the time interval between the Orrorin femur and the unpublished Aramis skeleton is longer than the time between Aramis and Hadar. Those relative times may become quite important in thinking about the evolution of those postcrania.

    The Winter Line was broken at Monte Cassino, after many failed attempts from different approaches. The Aramis fossils are either the heavy shoe waiting to drop, or they are the uncomfortable foot that all this talk about phyletic evolution is meant to shoehorn into place.

    Commentary

    If all these cases are added together, they imply a single evolving lineage encompassing at least four anagenetic taxa, Ar. kadabba -- Ar. ramidus -- Au. anamensis -- Au. afarensis. This last would presumably be followed by a cladogenesis into a robust australopithecine species (Australopithecus aethiopicus) and Australopithecus africanus.

    One could add Homo erectus to this list, since White and colleagues argued in their description of the Daka skull (Asfaw et al. 2002) that the Asian and African samples represent one cosmopolitan species.

    But then one species sticks out as a surprising exception to the pattern: Australopithecus garhi (Asfaw et al. 1999). It will be interesting to see a close argument showing why this species is really different from South African Au. africanus. Say, more different than KNM-WT 40000 is from the Hadar crania. It's quite glaring, really, that this species should be there mucking up such a simple phylogeny.

    I have to say, after reviewing all these papers in one sitting -- this entire bush vs. ladder thing is getting very tiresome! I mean, isn't there something else that we could organize early hominid discoveries by? These are all papers in the top journals, and this is the (fairly specialized) discussion that has been promoted as the central issue in the field!

    The subtitle of the Dalton piece suggests that it is merely a philosophical difference:

    Deciding whether our ancestors evolved as a single lineage may depend more on philosophy than fossils.

    But that's not really true. There is a clear null hypothesis here, quite directly drawn from William of Ockham:

    entia non sunt multiplicanda praeter necessitatem

    Which of course means:

    Sometimes fossil samples really do form ancestor-descendant relationships.*

    (*) It doesn't really. It means "Entities should not be multiplied beyond necessity."

    References:

    Asfaw B, Gilbert WH, Beyene Y, Hart WK, Renne PR, WoldeGabriel G, Vrba ES, White TD. 2002. Remains of Homo erectus from Bouri, Middle Awash, Ethiopia. Nature 416:317-320. DOI link

    Asfaw B, White T, Lovejoy O, Latimer B, Simpson S, Suwa G. 1999. Australopithecus garhi: A new species of early hominid from Ethiopia. Science 284:629-635. DOI link

    Begun DR. 2004. The earliest hominins -- is less more? Science 202:1478-1480. DOI link

    Brunet M. and 37 others. 2002. A new hominid from the Upper Miocene of Chad, Central Africa. Nature 418:145-151. DOI link

    Brunet M, Beauvillain A, Coppens Y, Heintz E, Moutaye AHE, Pilbeam D. 1995. The first australopithecine 2,500 kilometres west of the Rift Valley (Chad). Nature 378:273-275. DOI link

    Dalton R. 2006. Feel it in your bones. Nature 440:1100-1101. DOI link

    Haile-Selassie Y. 2001. Late Miocene hominids from the Middle Awash, Ethiopia. Nature 412:178-181. DOI link

    Haile-Selassie Y, Suwa G, White TD. 2004. Late Miocene teeth from Middle Awash, Ethiopia, and early hominid dental evolution. Science 303:1503-1505. DOI link

    Leakey MG, Spoor F, Brown FH, Gathogo PN, Kiarie C, Leakey LN, McDougall I. 2001. New hominin genus from eastern Africa shows diverse middle Pliocene lineages. Nature 410:433-440. DOI link

    Ohman JC, Lovejoy CO, White TD. 2005. Questions about the Orrorin femur. Science 307:845. DOI link

    Senut B, Pickford M, Gommery D, Mein P, Cheboi K, Coppens Y. 2001. First hominid from the Miocene (Lukeino formation, Kenya). Comptes Rendus 332:137-144.

    White T. 2003. Early hominids -- diversity or distortion? Science 299:1994-1996. DOI link

  • Earliest stone tools on Java

    Mon, 2006-04-24 13:24 -- John Hawks

    The current Science has meetings reports from the Indo-Pacific Prehistory Association Congress, including this article by Richard Stone about excavations from Sangiran:

    At the meeting, archaeologist Harry Widianto of the National Research Centre of Archaeology in Yogyakarta, Indonesia, wowed colleagues with slides showing stone tools found in sediments that he says were laid down 1.2 million years ago and could be as old as 1.6 million years. The find, at a famous hominid site called Sangiran in the Solo Basin of Central Java, "opens up a whole new window into the lifeways of Java Man," says paleoanthropologist Russell L. Ciochon of the University of Iowa in Iowa City.

    This report is the latest in a series of finds from the site, which have been increasing in date:

    Over 2 months, they unearthed 220 flakes--several centimeters long, primarily made of chalcedony, and ranging in color from beige to blood red--in a 3-by-3-meter section of sand deposited by an ancient river.

    The find, not yet published, could be even more spectacular than Widianto realizes, says Ciochon. His team, which also works at Sangiran, has used ultraprecise argon-argon radiometric methods to date the volcanic strata overlying the levels excavated by Widianto to 1.58 million to 1.51 million years ago--making the flakes at least 1.6 million years old. If the flakes were undisturbed, Ciochon says, they would represent "some of the earliest evidence of the human manufacture of stone artifacts outside of Africa." Their antiquity would match that of the oldest flakes found in China, at Majuangou, dated to 1.66 million years ago and also made of chert.

    The article includes a picture of some of the tools, which are small flakes on a fine-textured material.

    The Sangiran flakes "are fundamentally different"--smaller--than the stone choppers made by H. erectus in Africa, says Ciochon. The evidence, he argues, suggests that Java Man had to range far for small deposits of good flint or chert and so created small, finely worked tools in contrast to the larger tools found in Africa.

    There certainly is a contrast in the absence of large core tools, but it would take a close study to show that the flakes are different. But if there was a conscious limit to the selection of good raw materials, that might be significant also.

  • Lumbar vertebrae in hominids: six or five?

    Sun, 2006-04-23 21:42 -- John Hawks

    So have we decided that early hominids had five lumbar vertebrae?

    I ask because I was reading a paper by Owen Lovejoy (2005) the other day, and came across this passage:

    While most observers have concluded that both STS-14 and STW-431 exhibit six lumbar vertebrae, Haeusler et al. recently presented detailed arguments that these two specimens exhibit only five lumbar vertebrae (as well as KNM-WT 15000, a specimen of H. erectus also
    described as having six lumbar vertebrae; see below), but with the last thoracic essentially having lumbar-like function. Their arguments are based on traditional anatomical analysis (presence/absence and location of rib facets, etc.), and need not be considered further here. Even they conclude that all "three fossils differ from the majority of modern humans in that it is the seventh presacral vertebra where the orientation of the articular facets changes" (Haeusler et al. 2002, p. 634), thus making lordosis possible over a longer range of the lower back (Lovejoy 2005:102).

    There are two questions here: first, how many lumbar vertebrae (5 or 6) did early hominids have, and second, how many vertebrae (whether lumbar or thoracic) did they have functioning as part of the lumbar curve?

    Haeusler addresses the first question with anatomical examination of the three most complete vertebral columns from early hominids: Sts 14, STW 431, and KNM-ER 15000. From the abstract:

    Thus, in Sts 14 the sixth last presacral vertebra has on one side a movable rib. In Stw 431, the corresponding vertebra shows indications for a rib facet. In KNM-WT, 15000 the same element is very fragmentary, but the neighbouring vertebrae do not support the view that it is L1. Although in all three fossils the transitional vertebra at which the articular facets change orientation seems to be at Th11, this is equal to a large percentage of modern humans (Haeusler et al. 2002:621).

    The problem in all three cases is that the incompleteness of the column allows doubt about the number of thoracic vertebrae. Their argument is based on the morphology of the putative L 1 / T 12 in Sts 14 and STW 431. The most difficult case seems to be KNM-WT 15000 (the Nariokotome skeleton). Haeusler et al. (2002) discuss the critical differences as follows:

    The principal difference between Brown et al.'s (1985) interpretation of five and Walker and Leakey's (1993) six lumbar vertebrae presumably rests on the twelfth thoracic vertebra that the latter authors reported to be missing. This assumption might be based on the impression of an abrupt increase in the sagittal corpus height from vertebra Y to AR/BA (cf. Figure 7). Metrical dimensions, however, do not show abrupt changes between these two segments and hence do not oppose a position of vertebra Y immediately cranial to AR/BA (Figure 8). Therefore, Walker and Leakey's (1993) inference that a vertebra is missing at the thoraco-lumbar border is questionable.

    On the other hand, due to its poor preservation, the morphology does not allowidentification of vertebra AR/BA either as the first lumbar element (cf. Walker and Leakey, 1993) or as Th 12, as preferred here. The dorsal part of the body together with the pedicles is missing and thus it is unknown whether or not this vertebra had costal facets. Moreover, almost nothing is preserved of the processus spinosus. The sequence in the morphologies of the processus spinosi from the caudally adjacent vertebra AA/AV ( = L 2 according to Walker & Leakey, 1993) to that of vertebra Y ( = Th 11), however, makes it rather unlikely that a further -- not preserved -- vertebra came in between them in addition to AR/BA. This suggests that AR/BA, in fact, represents Th 12 instead of L 1, and AA/AV would become L 1 instead of L 2. The position of the anticlinal vertebra seems to support this hypothesis. In the modern human spine it is typically at the level of Th 11, with a range from Th 10 to Th 12 (Double, 1912; Danforth, 1930; Ankel, 1967). In KNM-WT 15000, the processus spinosus of Th 11 still projects caudally, which means that Th 12 must have been the anticlinal vertebra. On the other hand, the morphologies of the processus spinosi of vertebra Y and AA/AV -- though fragmentary -- are not far from that of an anticlinal vertebra. This again suggests that AA/AV represents L 1, and thus AR/BA becomes Th 12 (Haeusler et al. 2002:632-633).

    Ultimately, Haeusler et al. (2002) and Lovejoy (2005) come down to the same point: if you want to figure out how many of which type of vertebrae there are, then you need to know how many vertebrae there are, and none of the early hominid vertebral columns are complete. It would be easier to say that the last thoracic was actually the first lumbar if you knew that it would be T 13 instead of T 12.

    From that perspective, Lovejoy's argument really gains traction concerning the function of the putative first lumbar in the lordosis. This is important to his interpretation of the lower back morphology of apes:

    The implications are quite profound. Since apes exhibit the opposite change of lumbar column reduction (Fig. 2), the demonstrably more lordotic column in Australopithecus than occurs even in most H. sapiens signals skeletal reorganization assuring virtually complete abandonment of arboreal activity in favor of permanently terrestrial life. That is, if climbing induces the entire elimination of any mobility in the lower backs in (essentially quadramanual) arboreal apes, it is, by extension, antithetical to arboreality in a species in which so many other critically important adaptations for climbing had also been abandoned (see below) (Lovejoy 2005:102).

    Lovejoy traces the functional necessity for lumbar reduction in apes to the reduction in the m. erector spinae:

    Apes can only have afforded such reduction of their erector spinae if it was simultaneously accompanied by "passive" spinal immobilization so as to prevent habitual Euler buckling of the column (that is, by virtue of the combined deformation of its individual disks), though shortening of the column has largely eliminated this prospect in apes. Such changes were very likely also associated with other modifications of the thorax (and abdomen), including an "imagination" of the spine ventrally into the thorax (Fig. 6), a character which increases column rigidity and which was presumably associated with their more posterolaterally repositioned shoulder girdle than occurs in monkeys (ibid.).

    Well, I don't know if the five lumbar explanation is quite baked in yet, but it might take another vertebral column to turn it around.

    References:

    Lovejoy CO. 2005. The natural history of human gait and posture Part 1. Spine and pelvis. Gait Posture 21:95-112. DOI link

    Haeusler M, Martelli SA, Boeni T. 2002. Vertebrae numbers of the early hominid lumbar spine. J Hum Evol 43:621-643. DOI link

  • Acheulean endings

    Tue, 2006-03-28 22:57 -- John Hawks

    There is no hard endpoint to the Acheulean; its tool types -- in particular the handaxe -- last well into the MSA/Middle Paleolithic. Here are some notes on the later occurrences in Africa:

    Sheppard and Kleindienst (1996) considered the technological changes from ESA to MSA at Kalambo Falls, finding:

    ...there is little change, at this site, in the basic techniques of blank production or the attributes of the blanks produced from the ESA to the MSA. The only marked change to occur is the loss of large cutting tools (hand axes, cleavers) and their replacement by heavy-duty forms (core axes, picks). It is hypothesized that this change marks a decline in portability as a factor in the design of large edge tools (Sheppard and Kleindienst 1996:171).

    McBrearty (2003) reflects on the late persistence of characteristic Acheulean handaxes in association with some of the earliest fossils of "modern" humans, the Herto hominids from the Middle Awash of Ethiopia:

    At 160-154 ka, the Herto handaxes are the latest survivors of Acheulean technology known in Africa. Formerly, the best candidates were those from the site of Rooidam, South Africa, where a U-series date indicated they might date to ~ 170 ka (Szabo and Butzer 1979). K/Ar dates of ~ 240 ka for Acheulean artifacts from the Kapthurin Formation, Kenya (Leakey et al. 1969, Tallon 1978) have frequently been cited as the date for terminal Acheulean in east Africa, but new more precise 40Ar/39Ar dating now shows these to predate 285 ka (Deino and McBrearty 2002).

    What does the presence of Acheulean tools at 160 ka signify? The nature of archaeological change in the African Middle Pleistocene is murky, but the most obvious development is the abandonment of Acheulean technology, usually thought to have been made by H. erectus and its replacement by implements of MSA traditions, believed to have been produced by H. sapiens. In a nutshell, this comprises the replacement of handaxes and cleavers by points, signifying a shift from hand-held to hafted tools, and the birth of projectile technology. The presence of Levallois technology is usually considered to be a feature of the MSA, but it is present in late Acheulean contexts as well. In fact, the big flakes used to make African late Acheulean handaxes, including some Herto and Kapthurin specimens, may be struck from Levallois cores....The earliest MSA points were once thought to be those from Gademotta, Ethiopia, dated by K/Ar to 235 ka (Wendorf et al. 1994). Retouched points from the Kapthurin Formation predate those from Gademotta by 50,000 years. Dated by 40/Ar/39Ar to > 285 ka (Deino and McBrearty 2002), the are currently the oldest in Africa. A similar date is estimated for the basal MSA at Florisbad, South Africa (Grün et al. 1996), though these levels have not yet yielded points (Kuman and Clarke 1986, Kuman et al. 1999). Both retouched points and Levallois points have been recovered at Twin Rivers, Zambia, where they are dated by U-series to 265 ka (Barham 2000) (McBrearty 2003:1-2).

    McBrearty briefly discusses two alterative hypotheses. One is that the Acheulean handaxes are markers of a biological species, and that MSA points and other artifacts were made by some other species, or multiple species. The problems with this hypothesis are manifold, including the association of the handaxes with fossil remains that would be uncomfortable fits in H. erectus at best. Yet, the hypothesis can be sustained by the additional assumption that multiple species cooccupied the same African paleolandscapes with sufficient contemporaneity to drop their tools in the same places:

    Or can the archaeological record of the African Middle Pleistocene represent the behaviour of several species? Might different tools be the distinctive signatures of different populations or species who are competing with each other for the same territory and resources? A similar scenario of interspecific competition seems to have been played out between H. sapiens and the Neanderthals in Europe after 40 ka, but there the replacement was complete within 10,000 years or so. Africa is a far more enormous continent than Europe, its Pleistocene population far less dense, and our knowledge of its record less finely resolved. The 125,000 years represented by the Acheulean to Middle Stone Age transition in Africa is more than four times greater than the entire timespan of the residence of H. sapiens in Europe, but only a handful of African sites from this long time period have yet been explored.

    The other hypothesis is that the handaxe appears wherever it is functionally relevant, regardless of phylogeny. Acheulean elements may have been independently invented again and again:

    Archaeologists routinely stress the uniformity of the Acheulean tradition, and rarely entertain the possibility that the category 'handaxe' is an artificial construct that we impose upon the material. Experimental work by Jones (1994) shows that the familiar tear-drop shape of the handaxe is a design compromise resulting from attempts to gain the maximum cutting edge from the minimum weight of stone. The same techniques of blank production and bifacial retouch were employed in many cases to make both Acheulean handaxes and MSA points, and in the Kapthurin Formation the two classes of artefacts can grade into each other. Furthermore the tear drop shape occurs repeatedly in the archaeological record in cases where there is no evidence of 'phylogenetic' relation. Many projectile points from Africa, Europe, the Near East, and even the New World have similar plan forms (McBrearty 2001; Otte 2003), but there is clearly no 'phylogenetic' relationship among them. Rather, the recurrence of this target form demonstrates the design constraints inherent in fractured stone, and it appears to have been rediscovered repeatedly by tool makers in the course of prehistory (McBrearty 2003:2-3).

    Of course, if the recurrence of the handaxe across the 1.4 million years of the Acheulean resulted from its function rather than a continuous chain cultural transmission, then it takes away one of the principal pieces of evidence that Acheulean people had any substantive culture at all.

    As far as handaxes are concerned, that may be a good thing. The maintenance of a single cultural tradition across much of three continents over a million years by exclusively social transmission seems incredible. Some have suggested that the handaxe is hardwired into the human genome, a proposition that seems even less credible (at least, to me). Absent these means of transmission, we are left with the proposition that the handaxe did not fade from the earth because of its functional utility -- either it was the tool that did the job the best, or it was the best tool that humans were capable of making that did the job adequately. If function and human ability combined to make it so persistent, it should not be the least bit surprising that the form should recur in later contexts -- in the scenario of recurrent invention, it never really went away. The production of handaxes on large Levallois flakes seems especially relevant.

    One view of the uniformity of handaxe variation across space is that humans maintained strong cultural links across large distances. The knowledge of handaxe production and the concept of handaxe shape were highly conserved cultural markers that were widely spread.

    But the alternative hypothesis, that handaxe form was repeatedly invented by hominids and actually served as the best possible tool given its purpose and hominid flaking capacities, does not require long-distance cultural interactions to maintain the uniformity of the tradition. Nor does it require the recurrent movement of people over vast distances to spread the information. Given the lack of evidence for such long-distance movement, this hypothesis gains a bit of credibility.

    Raw material utilization in the Acheulean is highly localized. People may have been using the large tools as repositories for further flaking, and they may have curated them for some time. But they did not transport them over vast distances. Ordinarily, Acheulean people do not appear to have transported material over distances greater than the possible daily movement of a single individual -- a few tens of kilometers.

    It raises an obvious question: Is high mobility (in other words, hunter-gatherer-like mobility) a shibboleth of the assumption that handaxe form required culture? Did earlier humans actually have very small, limited home ranges and daily movement patterns? After all, if humans carried handaxes around for their utility, and they didn't manufacture a new one every day, then some handaxes ought to represent week-long movements or even longer.

    References:

    Sheppard PJ, Kleindienst MR. 1996. Technological change in the Earlier and Middle Stone Age of Kalambo Falls (Zambia). Afr Archaeol Rev 13:171-196. DOI link

    McBrearty S. 2003. Patterns of technological change at the origin of Homo sapiens. Before Farming 3:1-6.

  • What won't a handaxe do?

    Sun, 2006-03-05 23:57 -- John Hawks

    I've had this working paper by Tony Baker on my desktop for awhile, and it has been discussed on some message boards. I wanted to link before I forget. It's a good online review of the problems interpreting Acheulean handaxes. Baker's preferred theory is that the "handaxes" were principally cores for flake extraction. This is not a novel view, but the unique aspect is the way he derives the argument from the physics of flaking:

    Homo erectus did not select small cores from which to extract flakes (make handaxes). I propose he chose large cores because he did not have the manual dexterity to externally support them and, therefore, he had to rely on the inertial support. He just let them lie flat in his hand or on his leg. This inertial support knapping meant all blows had to be directed toward the center of mass. Flakes scars could not pass the center of mass so the handaxe remained relatively thick. Additionally, with numerous flake removals the handaxe became smaller and more ovate or discoid in shape because the blows were being directed from the furthest edge from the center of mass toward the center of mass.

    This is all described well, which is what makes the paper worthwhile. This paragraph is a good encapsulation of the problem of the traditional view:

    The Lower Paleolithic researchers who believe the Acheulean handaxe was the desired product do not find their justification in its function, since its function is not understood (Bordes 1968:64; Debenath and Dibble 1994:130; Gowlett, Crompton and Yu 2001:612; Isaac 1977:12,144; McPherron 2000:73; Roe 1981:271). Instead, they find their justification in its unchanging morphology. For a million plus years, its basic shape remained constant as it spread across three continents. It has a tip, a butt, and often symmetry in three dimensions. Its length is rarely longer than twice its width, and Gowlett has demonstrated at Kilombe, that a linear relationship between length and width explains 69% (R=0.83) of the variation (1995:202). Issac proved there was a strong correlation between the length/width ratio and length regardless of sites or continents (1977:139). Further, its width is rarely greater than three times it thickness. This unchanging morphology is, therefore, proof that the handaxe was constructed to conform to an unchanging, deeply engrained mental template.

    When I looked at this first, I thought I would post an expanded review of the handaxe problem, but other matters delay that at the moment. In the meantime, Baker's paper is a valuable resource.

    In summary, my belief is that the handaxe was a large biface core made by Homo erectus individuals who lacked the manual dexterity of modern humans. I arrived at this conclusion based on the morphology of the handaxe. And, since there is no fossil hand evidence to suggest otherwise, my theory is intact. I am forced to admit that I believe Homo erectus' hand was evolving during its 1.6 million years of existence. Therefore, I would expect there also was an evolution in the handaxe from large to small and thick to thin. Yet, this evolution is only going to be visible in vertically stratified sites or well-dated sites. And, then it will still be difficult to identify because the handaxe was the by-product of expedient flake extraction. It was not the desired product.

    I think it would be better to replace "hands" here with "brains", as the prerequisite "dexterity" in this context probably is more mental than manual.

  • Tilting at absent Asian australopithecines

    Mon, 2006-01-09 00:27 -- John Hawks

    In Nature a couple of weeks ago, Robin Dennell and Wil Roebroeks had a provocative paper exploring the possibility that early humans (i.e. Homo erectus) originated in Asia rather than Africa.

    The paper is all speculation of course; there is no evidence of any earlier hominid in Asia.

    But it is the good kind of speculation. Although maybe not quite this big:

    Most probably, we are on the threshold of a profound transformation of our understanding of early hominin evolution that might prove as far-reaching as the demise of the notion of Man the Hunter in the early 1960s (Dennell and Roebroeks 2005:1103).

    Here's the abstract:

    The past decade has seen the Pliocene and Pleistocene fossil hominin record enriched by the addition of at least ten new taxa, including the Early Pleistocene, small-brained hominins from Dmanisi, Georgia, and the diminutive Late Pleistocene Homo floresiensis from Flores, Indonesia. At the same time, Asia's earliest hominin presence has been extended up to 1.8 Myr ago, hundreds of thousands of years earlier than previously envisaged. Nevertheless, the preferred explanation for the first appearance of hominins outside Africa has remained virtually unchanged. We show here that it is time to develop alternatives to one of palaeoanthropology's most basic paradigms: 'Out of Africa 1' (Dennell and Roebroeks 2005:1099).

    It is worth reviewing exactly what "Out of Africa 1" is supposed to be. The paradigm is that emergence of hominids from Africa required increases in brain size and/or body size, coincident with the emergence of hominids like KNM-ER 3733, KNM-WT 15000, and others. The motivation for this hypothesis is simple: australopithecines have not been found outside of Africa. Nor has anything like Homo habilis, which is australopithecine-sized but has larger brains.

    Of course, it is questionable just how basic this paradigm is. Consider what I (and my colleagues) were able to write only seven years ago:

    The problem is that significant range expansion out of Africa occurred a half million years or more later than the first H. sapiens [corresponding to others' H. erectus or H. ergaster]. Population size before then may have remained small, and this is not an inconsequential time span, being one quarter of the time H. sapiens has existed. An important date in behavioral evolution is 1.5 MYA because it is marked by the earliest appearance of the Acheulean, the ubiquitous hand-axe industry of the Early and Middle Pleistocene.... Before this time, humanity was limited to Africa and immediately adjacent sections of Asia such as the Levant (Hawks et al. 2000:7).

    Evidence for large body size in Late Pliocene humans (notably KNM-WT 15000 but also many others) made it very plausible that larger bodies were necessary for dispersal from Africa. But without good evidence for such dispersal before around 1.4 million years ago (and arguably not before 1 million years), larger bodies could not be assumed to be a sufficient condition for dispersal. Writing about the origin of humans, we had to consider all these alternatives -- at a time when the Dmanisi sample consisted of a single uncertainly dated mandible and the Mojokerto date stood alone with very questionable provenience.

    Now we know that hominids did leave Africa by at least 1.8 million years ago. Dmanisi has almost singlehandedly changed the perspective.

    And in doing so, it made much more convenient the hypothesis that large body size was both necessary and sufficient for dispersal from Africa. If the date of dispersal and the date of human origins are the same, then it is natural to propose that the coincidence is more than chance.

    I would say this is more of a convenient hypothesis (and an easy story to tell) than it is a basic paradigm. The idea that large body size caused dispersal from Africa may have been a local minimum in terms of parsimony (at least as long as the body size of the Dmanisi fossils was not known), but it was only one alternative among many still in play.

    And it remains a plausible hypothesis -- after all, the Dmanisi remains are a bit larger than australopithecines, and they might well have shrunk from a larger early-human-like size after reaching Asia instead of before.

    But Dennell and Roebroeks give motivations for examining some alternatives.

    The only reason why the earliest tool assemblages in Asia are attributed to H. erectus s.l. is that palaeoanthropologists have already decided that, in effect, it was the only hominin capable of migration out of Africa, and with sufficient Wanderlust to do so (Dennella and Roebroeks 2005:1099).

    Homo erectus sensu lato (s.l.) means Homo erectus "in the loose sense", which would include not only the "strict sense" (sensu stricto) H. erectus. from Java and China, but also hominids like OH 9 and KNM-ER 3733 from Africa, and presumably the Dmanisi hominids.

    A long passage reviews the total faunal evidence from Asia during the Late Pliocene. The thrust of the passage is that there are very few sites with extensive fauna, and of these most preserve mainly large-bodied herbivores. There are a few hints that a hominid-friendly fauna may have existed, including the presence of baboons. But there are no hominids of any kind at the vast majority of Asian localities -- Dmanisi is a real exception in the Plio-Pleistocene record.

    This is the key taphonomic argument: if we have only found Early Pleistocene humans from continental Asia within the past ten years, then how can we preclude there having been australopithecines there? Dennell and Roebroeks argue that if there were australopithecines, we shouldn't necessarily expect to have found them yet -- we just haven't looked extensively enough.

    A close read of the section raises a caution, though. One of the main arguments for the incompleteness of the Asian record is that sites don't preserve each others' fauna.

    It is also likely that the full range of taxa is incomplete for the Indian subcontinent, because Megantereon and Pachycrocuta are not recorded in India but are present in Pakistan; in Pakistan, there is no evidence of Camelus and small primates, and in neither country is Homotherium recorded, although this is present to the west at Dmanisi, to the north at Kuruksay, central Asia and to the east at Longuppo, south China (Dennell and Roebroeks 2005:1100).

    Of course, all of these species are recorded in Asia taking all the sites in aggregate; this is hardly an argument for the overall weakness of the record -- just an argument that no individual site is an adequate record of the continent's fauna.

    To me, the important question is not whether australopithecines as currently known from Africa were in Asia. A more troubling possibility is that the australopithecines that we now know from Africa were not the only (or main) manifestations of early hominids in Africa. Large parts of Africa that we might expect to be congenial to hominids, like the Zambesi basin, have few or no fossils at all. The recovery of the Bahr el Ghazal mandible (Brunet et al. 1994) certainly makes clear that hominids were living across a much larger area than we have adequately sampled. But that mandible is, although not identical, certainly very similar to known contemporary hominids in its adaptation.

    The question is whether hominids had adapted to other ecologies that are much less satisfactorily sampled than the East African rift. They probably weren't living where chimpanzee and gorilla ancestors did, but where else might they have been? Some such ecologies -- like the coasts -- would make early dispersal very plausible.

    (In this regard, early humans are not the only hominids who lack a satisfactory ancestor. Who was the ancestor of A. aethiopicus? In what ecology did the first robust hominid arise?)

    So what is the broader set of hypotheses that we should consider? Dennell and Roebroeks suggest:

    If the above taphonomic review suggests that we cannot show the absence of hominins from areas in Asia at a time before the little evidence we have indicates their presence, we need to consider alternatives to the current Out of Africa [that is, their "Out of Africa 1"] model. There are three issues here. The first is when hominin(s) first left Africa -- might they, for example, have left shortly after they acquired the ability to make stone tools, the earliest of which are currently 2.6 Myr old? Or could they have left even earlier, about 3.0Ð3.5 Myr ago, when some australopithecines were already living in the African grasslands? The second issue is whether we yet know the full range of hominins that inhabited both Africa and Asia in the Late Pliocene and Early Pleistocene. Even in east Africa, several new taxa have been claimed in the past decade (for example, A. anamensis, A. garhi, Ardipithecus ramidus and Kenyanthropus platyops) and doubtless more will be found. (An indication of how little we know about Pleistocene east Africa is that only recently has the first fossil evidence for chimpanzee been found.) In Asia, the recent discoveries of H. georgicus and H. floresiensis should make us very wary of assuming that H. erectus s.l. was the only player on the Asian stage in the Early Pleistocene. Third, Asia might not have been the passive recipient of whatever migrated out of Africa but might have been a major donor to speciation events, as well as dispersals back into Africa. Such two-way traffic is well documented for other mammals in the Pliocene and Early Pleistocene, such as Equus and bovids, with more taxa migrating into than out of Africa. There is no reason why hominin migrations were always from Africa into Asia, and movements in the opposite direction might also have occurred, as has been suggested for the Olduvai OH9 (refs 13, 58) and Daka specimens. We should even allow for the possibility that H. ergaster originated in Asia and perhaps explain its lack of an obvious east African ancestry as the result of immigration rather than a short (and undocumented) process of anagenetic (in situ) evolution (Dennell and Roebroeks 2005:1100-1101).

    Of course, most of the evidence indicating the presence of hominids is not fossil but archaeological. On this topic, Dennell and Roebroeks have much to say:

    Any stone tool assemblage in Asia dated as older than 1.9 Myr ago (the earliest date that Homo is supposed to have left Africa) is either dismissed or (more usually) ignored; undated Oldowan tools are assumed to date from after 1.9 Myr ago and not from 2.6 Myr ago (the date of their first appearance in east Africa); and stone tool assemblages in Asia dated to the Olduvai Event (1.77Ð1.95 Myr ago) and not associated with hominin remains are automatically attributed to Homo erectus s.l. However, there is no reason why Oldowan assemblages in Arabia cannot be older than 1.9 Myr old, or why the tools from Ain Hanech (Algeria) or Erq el Ahmar (Israel) were made by H. erectus s.l. [instead of other hominids] (ibid:1102, references omitted).

    There is a section about what exactly absence of evidence can tell, a short critique of using continents as proxies for biogeographic units:

    As noted earlier, Pliocene grasslands extended all the way from west Africa to north China, and 'Savannahstan' might prove a more useful spatial unit for modelling early hominin adaptations and dispersals within them than simply an undifferentiated 'Africa' or 'Asia'. For example, the African hominins 1.9Ð1.7 Myr ago at Koobi Fora (Kenya) and Ain Hanech (Algeria), and their slightly later counterparts in Asia at 'Ubeidiya (Israel), and Majuangou (north China) were all living in broadly comparable grassland environments, and it makes sense to place them within the same frame of reference.

    I think there is much of value to consider here; but it is less a revolution and more a statement of the field in transition. There are also alternatives that are not considered in this paper but that may be equally plausible -- most notably, the idea that early humans themselves may have been substantially polymorphic (witness KNM-ER 42700), or that brain size rather than body size may have been a prerequisite to dispersal (since habilines, Dmanisi, and H. erectus s.l. are all allometrically similar in brain size).

    National Geographic News also has an article about the paper.

    References:

    Dennell R, Roebroeks W. 2005. An Asian perspective on early human dispersal from Africa. Nature 438:1099-1104. Full text (subscription)

    Hawks J, Hunley K, Lee S-H, Wolpoff M. 2000. Population bottlenecks and Pleistocene human evolution. Mol Biol Evol 17:2-22.

  • Zimmer on Templeton

    Wed, 2005-12-07 10:12 -- John Hawks

    Carl Zimmer has a post discussing Alan Templeton's work. It's a good review, covering Templeton's two essential points: history cannot be traced from any single gene locus, and evidence from multiple loci makes human evolution look like a trellis of relationships among populations, not a diverging tree originating in one population. Here's a quote:

    Templeton has found that he can easily reject the idea that all our genes come from the same 200,000 year old population of African humans. Instead, he finds evidence of three separate expansions out of Africa. The first he estimates to have occurred 1.9 million years ago -- which just so happens to coincide with the earliest fossils of Homo erectus outside of Africa. Then he finds another expansion he dates to 650,000 years ago -- which just so happens to coincide with the emergence of hominids in Europe of hominids that are believed to give rise to Neanderthals. The last expansion can be traced back 130,000 years ago.

    I'll have more discussion here when Templeton's new paper comes out in the Yearbook of Physical Anthropology. In the meantime, you can get a quick review by reading the post.

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