john hawks weblog

paleoanthropology, genetics and evolution

Kenyanthropus

  • The trouble about Kenyanthropus and Ardi

    Thu, 2009-12-10 15:32 -- John Hawks

    There are three skulls from putative "hominins" that date to 3.5 million years or earlier. Every one of these skulls is known now from extensive reconstruction or correction for distortion in the original.

    By itself, the extensive reconstruction might not be a problem. But as Tim White has repeatedly shown, the specialists on these crania actively and vociferously disagree about the basic anatomy due to problems reconstructing them. White's ongoing dispute about the skull of KNM-WT 40000 is a matter of public record, both in his initial 2003 article on the skull, and in Michael Balter's description of the recent Royal Society meeting*:

    When the talk was thrown open for discussion, White took the microphone and began firing questions at Spoor about the degree of variation of the cheekbone position among specimens of A. afarensis and other hominin species. “We took that into account,” Spoor responded, “and I just showed you a graph” about it. “I didn’t ask you whether you took it into account; I asked you what it was,” White said. Spoor, clearly frustrated, told the audience that he had no vested interest in this debate. At that point, the session chair interrupted and invited everyone to break for coffee, but Spoor and White continued to debate between themselves for the next half-hour.

    If KNM-WT 40000 were the worst case, that would be bad enough. But Ardi's skull has required reconstruction even more extensive than would be required for the Kenyanthropus holotype.

    In their description of the Ardipithecus skull, Suwa and colleagues (2009) mainly present metrics taken from the CT reconstruction. The publication strikes me as remarkable in that it includes few photographs of the original fossil, and only one or two of the photos are in standard anatomical orientation. A substantial part of the CT reconstruction is based on a second individual (ARA-VP 1/500), of which no photographs are provided. For this, readers may refer back to the single rather poor photo in the 1994 description. Anatomical comparisons in the present paper are limited to visualizations of the CT reconstruction.

    As I've written elsewhere, I think that Suwa and colleagues did a remarkable piece of reconstruction. But it is non-replicable. The CT-reconstruction is a composite of two specimens that includes mirror-imaged parts. A tremendous amount of work went into it, but without access to the component parts, it isn't possible to test or verify the assumptions underlying the present model.

    Another striking thing about the Ardipithecus skull description is the lack of anatomical comparisons with relevant samples. I mentioned above that most of the figures involve metric comparisons -- many of them scaled to the cube root of endocranial volume -- which of course can only be taken on a small fraction of early hominin crania. That leaves out the most relevant specimens in the Hadar sample, including all the cranial specimens from AL 333. It leaves out most of the Sterkfontein collection.

    And it brings us back -- again! -- to Kenyanthropus. Reading back through the paper, it's hard for me to believe that reviewers allowed Suwa and colleagues to publish on Ardi's skull without including any comparisons with KNM-WT 40000. It's the earliest complete skull of an undoubted hominin.

    They're entitled to their opinion that the skull is distorted. I agree. But you can still compare most of its nonmetric features and put some reasonable bounds on its metrics. I mean, they included OH 5, for goodness sake -- which has nothing whatever to do with hominin origins. Including the comparisons wouldn't have changed much about the paper, although I'll point out that there's at least one derived feature of later hominins that KNM-WT 40000 and ARA-VP 1/500 both lack, and which isn't noted either by Suwa and colleagues (2009) or in the table presented by White and colleagues (2009).

    So what should we do? We can't see the scans, no independent reconstructions are possible, and the people who can see the scans refuse to present comparisons of these three skulls that together represent the supposed origin of the hominin lineage.

    AAAARRRRGGHHHH!

    We need to set up multiple sets of independent reconstructors having a replicable go at these skulls. These are the three earliest hominin skulls. Every one of them is crushed in some weird way. It would be a credit to the science to document their reconstruction in nauseating monograph-level detail. They're scans, for goodness' sake -- there's absolutely no argument that access should be limited for any reason.

    If I were running this, I would set up a graduate seminar devoted to putting them together, split among four universities, with results to be reported in a session at the meetings and monographically by e-publication. The issue is not whether we can obtain an exact representation of the original anatomy. The issue is whether we can reject hypotheses about that anatomy. Testing hypotheses requires us to survey the range of possible reconstructions and how they relate to the range of anatomical variation in living and extinct analogs. The more reconstructions, the better the testability.

    At the moment, that testability isn't there. I trust the anatomical expertise of the people who made the models, but they're just single models with no assessment of the range of error. I've written about the importance of open access for these reconstructions already ("Open access and fossil reconstruction"). The points here just amplify that theme.

    * As an aside, I wonder if the title of the Royal Society meeting ("The First 4 Million Years of Human Evolution") contemplated the possibility that there may have been only 4 million years of human evolution in total?

  • Why didn't they let Kenyanthropus save them?

    Thu, 2009-12-10 00:42 -- John Hawks

    In the fossil record, a species is a hypothesis. We can't test that hypothesis in the way we do with living animals. Even in the dark, after all the paleontologists have left, the fossil bones just won't get it on. No reproduction, no test.

    So, sometimes we have to live with hypotheses that we can't immediately test. Because many hypotheses are wrong, we have to keep juggling in our minds the names of more species than probably existed.

    All the juggling frequently leads to confusion. One may reasonably wonder how we know that species X evolved into species Y, when half the field rejects the hypothesis that X was a real species. Often we don't disagree about the "evolving into", but we do disagree about the boundaries and other relationships of the populations -- which we can understand only indirectly from the fossils. The fossils don't change, but our hypotheses about them do.

    That brings us back to A. anamensis. Here's a hypothesis about an ancient lineage of hominins, based on a certain number of differences from later fossil samples assigned to A. afarensis. That's a precarious place for a hypothesis, because from the beginning, the definition of A. afarensis has encompassed geographic and temporal variability. How hard would it be to recognize a little more temporal variability? Not very.

    So, as I described last week, Haile-Selassie and colleagues (2010) propose sinking A. anamensis ("Woranso-Mille: A ladder not a bush"). I should mention, if I haven't already, that I have great sympathy for this viewpoint. Absent some compelling evidence that the lineage includes a speciation event, I prefer slow gradual anagenesis to be categorized into one evolving species, not an arbitrary set of chronospecies.

    In the same post, I described the work of Kimbel and colleagues (2006), who had argued for anagenesis in the same sample of A. anamensis and A. afarensis-referred fossils, but retained the two distinct names for them. One thing stands out as a mystery to me in that paper. Why didn't they let Kenyanthropus make the argument for them?

    If you want to establish that A. anamensis is taxonomically valid, the simplest way to answer all critics is if it has more than one descendant. You don't have to demonstrate the phylogeny beyond all doubt, I would say, you just have to take the hypothesis seriously.

    In this instance, we seem to have two good candidates for a non-A. afarensis descendant of A. anamensis. The more obvious of them is Kenyanthropus. Why didn't they advance the hypothesis of a A. afarensis-Kenyanthropus clade? Here's what Kimbel and colleagues wrote about the latest Leakey find:

    A more significant concern is the presence of Kenyanthropus at 3.5 Ma. Kenyanthropus may demonstrate cladogenesis prior to this time, but this taxon is only directly relevant to the analysis if any of the samples share derived character-states with it. At present, the Kenyanthropus hypodigm does not match the others in the availability or quality of character data for the mandible and anterior teeth, while the evidence that exists (from the maxilla, for example) does not suggest a close relationship of Kenyanthropus to any of the phena considered here (Leakey et al., 2001).

    So, they took the hypothesis off the table. And again, they've refocused the question solely upon dental and mandibular evidence.

    This is a very large hole. At 3.5 million years, KNM-WT 40000 is earlier than any other comparably complete hominin skull, except for Ardi and Toumaï. It doesn't look like them, that's for sure. Not the same phenum at all. If you're going to insist that KNM-WT 40000 isn't A. afarensis, it's hard to see a better hypothesis than that it's descended from A. anamensis.

    If it doesn't have a close relationship to either A. afarensis or A. anamensis, then I'm at a loss to figure out what they think it is related to!

    I'm willing to believe White's (2003) argument that it just is a member of the A. anamensis-A. afarensis lineage, but I don't see the contrary argument that it's so different from this lineage that it must be derived from some as-yet-undiscovered hominin. Keep in mind that the argument was formed by people, many of whom already knew basically what Ardi's skull was going to look like. I just can't feature why this phylogenetic problem didn't raise itself to a higher profile.

    As I wrote above, we juggle more hypotheses than can be true. In this instance, the null hypothesis is that all these hominins belong to a single evolving species, which would be called A. afarensis. But one alternative, in which A. anamensis existed as the ancestor of A. afarensis and at least one other species, has some utility. It lets us refer clearly to phylogenies with late-diverging sister taxa. As Yoda might have said, there is another possible sister taxon for A. afarensis: A. africanus might be derived from, or might itself be, a South African contemporary of the Hadar-Maka-Laetoli sample. The earliest Sterkfontein dates go up and down; one or more of the remains might be contemporaries of Laetoli or even earlier East African localities.

    Anyway, I'm not so interested in this question of bushes versus ladders, or "Pliocene diversity". I'm more focused on the curiously non-hominin-like features of Ardipithecus. If we suppose a late molecular divergence of hominins and chimpanzees (say after 4.4 million years ago), then Ardipithecus might be an ape or ancestral (stem) hominid, not a hominin. If so, then samples now referred to A. anamensis, including later specimens from Aramis, Asa Issie, and Kanapoi, are in fact the earliest-known members of the human lineage.

    A. anamensis is no afterthought in that case, it may be the stem hominin.

    That is, if it hasn't been sunk into A. afarensis.

    UPDATE (2009-12-10): A reader writes to remind me about A. garhi, which presents itself as very much like A. africanus and is quite a lot later than any known A. afarensis samples. One alternative is that A. africanus is simply the latest element of the single A. anamensis-A. afarensis-A. africanus anagenetic sequence.

    I don't think that's very easy to reject, considering the lack of good cranial remains between 3 and 2.5 million years ago (or for that matter, even later) in East Africa.

    Some of the Sterkfontein specimens, particularly those from Jakovec cavern and from Member 2 (including Little Foot) have been suggested to be older than 3 million years. Partridge and colleagues (2003) put them at older than 4 million years ago, which would make them rivals of A. anamensis as the earliest hominins. But Berger and colleagues (2002) argued (apparently preemptively) that these early dates are not necessary on faunal or magnetostratigraphic grounds.

    A second reader wonders why I didn't mention A. bahrelghazali as a possible sister to A. afarensis. Well, nothing's impossible, but I'd say the case for the Bahr el Ghazal mandible being distinct from the Hadar-Maka-Laetoli sample isn't very strong. Still, we're only talking about hypotheses here, I suppose.

    References:

    Berger LR, Lacruz R, de Ruiter DJ. 2002. Revised age estimates of Australoipithecus-bearing deposits at Sterkfontein, South Africa. Am J Phys Anthropol 119:192-197. doi:10.1002/ajpa.10156

    Haile-Selassie Y, Saylor BZ, Deino A, Alene M, Latimer BM. 2010. New hominid fossils from Woranso-Mille (Central Afar, Ethiopia) and taxonomy of early Australopithecus. Am J Phys Anthropol (in press) doi:10.1002/ajpa.21159

    Kimbel WH, Lockwood CA, Ward CV, Leakey MG, Rak Y, Johanson DC. 2006. Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record. J Hum Evol 51:134-152. doi:10.1016/j.jhevol.2006.02.003

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

    Suwa G, Asfaw B, Kono RT, Kubo D, Lovejoy CO, White TD. 2009. The Ardipithecus ramidus skull and its implications for hominid origins. Science 326:68e1-68e7. doi:10.1126/science.1175825

    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.

    Partridge TC, Granger DE, Caffee MW, Clarke RJ. 2003. Lower Pliocene hominid remains from Sterkfontein. Science 300:607-612.

    Synopsis: 
    If A. anamensis evolved into A. afarensis, why do we need two names for them?
  • The teeth that didn't bark

    Sat, 2009-12-05 15:00 -- John Hawks

    Earlier in the week, I wrote about the new interpretation of fossil teeth from Woranso-Mille, Ethiopia ("Woranso-Mille: A ladder not a bush"). There was one aspect of the paper that I wanted to comment at some greater length: Why didn't the paper include comparisons with the Lomekwi sample of teeth and mandibles?

    The Lomekwi sample includes more teeth than the sample reported from Woranso-Mille. At around 3.5 million years old, Lomekwi is a more relevant comparison than Hadar or Kanapoi, closer in age to Woranso-Mille than any sample other than Laetoli. So why didn't they include the comparison?

    The obvious answer is that these teeth belong to Kenyanthropus platyops, a different species off the line leading from A. anamensis to A. afarensis. So they're not relevant to testing hypotheses about evolution within that lineage.

    But...I don't see why that answer is obvious if Kenyanthropus is a fictitious species, based on a distorted skull (e.g., White 2003).

    Suppose that the teeth don't represent Kenyanthropus. Then they ought to belong to the one cosmopolitan species that exists both north and south of Lomekwi, both earlier and later in time. They ought to be A. afarensis. Which makes them directly relevant to the Woranso-Mille hominins. At the very least, they add to the variation of the A. afarensis sample, helping to inform about the temporal trend in that lineage.

    Besides that, even if we thought that Kenyanthropus was a real taxon, most of the Lomekwi teeth might still be A. afarensis. Only two specimens were assigned to K. platyops by Leakey and colleagues (2001): the KNM-WT 40000 skull and the unassociated maxilla fragment KNM-WT 38350. Both those have relatively small molars compared to the known A. afarensis sample, but in both cases the teeth are broken and dimensions are estimated. The rest of the dental sample is unassociated. The only thing keeping them from being A. afarensis is what we're willing to assume about the number of species at Lomekwi.

    So, it seems like a comparison that ought to be done. The information content is not going to be really high -- we're only talking about gross dimensions of the crowns, and the only ones that look informative at all are upper molars. But Leakey and colleagues argued that the KNM-WT 40000 and KNM-WT 38500 molars are at the very lower end of the range of A. afarensis. Guess what? The Woranso-Mille sample extends this range downward, right around the Lomekwi range.

    Something there...

    UPDATE (2009-12-08): I should add that Brown and colleagues (2001) reported additional teeth from Lomekwi and other localities in the Turkana Basin, which they referred to A. afarensis on the basis of their temporal position. They're not in the Woranso-Mille comparisons either.

    References:

    Haile-Selassie Y, Saylor BZ, Deino A, Alene M, Latimer BM. 2010. New hominid fossils from Woranso-Mille (Central Afar, Ethiopia) and taxonomy of early Australopithecus. Am J Phys Anthropol (in press) doi:10.1002/ajpa.21159

    Kimbel WH, Lockwood CA, Ward CV, Leakey MG, Rak Y, Johanson DC. 2006. Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record. J Hum Evol 51:134-152. doi:10.1016/j.jhevol.2006.02.003

    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.

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

  • Woronso-Mille: A ladder not a bush

    Tue, 2009-12-01 02:58 -- John Hawks

    In a new paper, Yohannes Haile-Selassie and colleagues describe new hominin fossils from Woranso-Mille, Ethiopia. A good thing: It gives somebody like me a rationale for describing early hominins from the point of view of Hadar. You see, Hadar is the first sample to include a really complete skeletal representation. You can present earlier sites as a series of "firsts", but that's kind of misleading. Now, the simple Ardi-Lucy comparison carries a lot of water for teaching early hominins, and if we can assume that the samples intermediate in time are mostly A. afarensis-like, so much simpler.

    Oh, and one more really good thing: Standard dental measurements are provided in the text of the paper. Thank you, AJPA! We may not get all the specimens, but at least we can check the statistics.

    But there's a chance that things are not so simple as they seem, that there are mysteries still waiting to jump out of this sample and scare us at night. I imagine that some people are less than thrilled about this paper, which explicitly rejects the reality of one Leakey-named species and ignores another into obscurity. One might expect me to welcome our new lumping taxonomic overlords. And yet, this little paper doesn't provide some information and comparisons that seem like curious omissions. Which makes me wonder...

    The fossils from Woranso-Mille are between 3.6 and 3.9 million years old -- basically older than Laetoli and younger than Kanapoi. Since the Laetoli sample is A. afarensis, and the Kanapoi sample is A. anamensis, we can expect that the Woranso-Mille sample would say something about how these two species were related to each other. The fossils might be one species or the other, they might be intermediate between them, or they might even be something altogether different.

    What is there?

    The sample as described is almost exclusively dental, with only a fragment of mandible and another of maxilla tossed in the mix.

    Some readers may have been under the impression there's more at this site, and indeed I am as well. I think I've even seen them for 500 milliseconds at a meeting once upon a time. Of course, maybe that was a dream. Much in paleoanthropology seems to be fading into a unicorn fairyland these days...

    Wait a minute! It's for occasions like this that I have a blog! As it turns out, I took some notes on Woranso-Mille back in 2007.

    Now, I have to warn you: These notes were so snarky that I didn't dare hit "publish". But there's no sense shirking responsibility for them now. Next thing I know, some crank will be hacking my server to bring all this snark into the open!

    Along with many other people, I got to see the hominids from Woronso-Mille this spring. Then again, see is probably an overstatement. I mean, when you see something, generally light waves from the objects actually have time to strike your retinas. I couldn't swear that anyone actually had that experience during Yohannes Haile-Selassie's talk to the Paleoanthropology Society. Sure, there was a subliminal impression that the pictures were there. And yet, Powerpoint and automatic timing can do magical things.

    I experimented a bit later, to try to estimate just how long the pictures had been up there. The 500 millisecond setting seemed about right. Definitely automated. Too short for microsaccades to bring in the edges of the fossils properly. And many of them were in situ photos, with a lot of brown-on-brown. Hard to pick out edges at all, and some edges were still in the ground.

    I mean, really, work out the time that Santa Claus has to spend in each kid's house on Christmas Eve to make it to all the world's children in one night. That's the kind of time we're talking about.

    See what I mean. I mean, that's over-the-top snark. Still, it's better material than I usually work with, so I can't for the life of me figure out why I didn't publish it. It goes on:

    Don't get me wrong. I think it's entirely appropriate to hide the images, dim them, heck, don't even show them if they don't want to. Think of all the yokels like me who could tell immediately from a decent picture whether the fossils were A. afarensis or not, and go shooting off their servers to the rest of the world. Hard work in the field, with the high risk of failure, deserves every possible reward -- certainly the right to take the necessary time to make a careful analysis. I hardly ever make any comments after I hear a public talk, unless the material is already well-known or described elsewhere. And there are other practical reasons not to talk about it -- for one thing, people sometimes change their minds!

    But why should I feel any compunction about prognosticating on fossils that are announced in the press? Hey, if they didn't want the attention, they wouldn't have a press conference, right? I'll bet they didn't make the press sit through the half-second slide show!

    Haile-Selassie announced several of the Mille fossils in 2005, notably the partial skeleton -- of which they are still trying to find more parts. At the time, I wrote about it, Rex Dalton wrote about it, Ann Gibbons wrote about it, twice, the AP wrote about it. Good times were had. Oh, those good times. Sure, no descriptions. Granted, in situ brown-on-brown photographs with buried edges. But good, good times.

    How could I have forgotten those good times?

    Now, there is a second press offensive underway. The best stories are at National Geographic News and The Cleveland Plain Dealer (Haile-Selassie's in Cleveland). It's an important site, with dozens of specimens.

    Hey, maybe they're like the Laetoli footprints and they rebury them when they're done looking at them. Kind of like catch-and-release.

    The stealth mandible

    All this was nearly three years ago. Which is a bit strange, considering that the current paper still doesn't include all the specimens. Assuming the 2007 illustrations were correct, the current paper doesn't even include all the Woranso-Mille dental specimens, as at least one mandibular dentition appears to have been omitted. It is, of course, possible that the news reports had the wrong picture.

    Here are my 2007 thoughts on the matter:

    We can probably answer this already: the National Geographic story includes a picture of the most complete mandible, and it looks an awful lot like LH 4, maybe with a more sloping symphysis. It's a rotten view - artistic, sure, but a lousy angle for comparison.

    This mandible is not included in the current paper. It is pretty obviously the most diagnostic of the mandibular/dental specimens, if it's from Woronso-Mille. I wonder if National Geographic really may have credited the wrong photo to Haile-Selassie? Very strange. In any event, it's an important question since the sample of other postcanine teeth in the paper is generally 2-3 specimens. A missing postcanine dentition would make a lot of difference to our picture of the variation.

    OK, continuing on:

    But still, the teeth appear to fall into the Laetoli-Maka-Hadar sample, the postcanine rows diverge from each other, and the symphyseal morphology in A. afarensis is certainly variable enough to encompass this mandible. Really the only missing feature that would be helpful is the P3, but unless other specimens turn out to be outside the Hadar range, I would assume this is going to be assigned to A. afarensis.

    Which does make me wonder how much the hidden mandible has driven the paper's conclusion. On the basis of the specimens they published, the majority of dental features seem to argue in favor of A. anamensis, as they explicitly write. They mention only a few "derived features" also present in later sites. Given the date, one might just as easily argue that these "derived" features were actually low-frequency variants heretofore unrecognized in the small A. anamensis sample, so that Woranso-Mille extends the range in that species while maintaining its overall anatomical pattern. The stealth mandible, if indeed it exists, looks more persuasive fitting in the pattern of A. afarensis.

    The unanswered phylogenetic questions are chiefly about what other lineages there may have been at the time. Mille might answer that question if substantial hominid diversity were found there, or at least something really different from the other sites. But no apparent evidence of such diversity was apparent in the public lecture. Maybe there are surprises waiting, but this team in the past has argued pretty strongly for taxonomic conservatism.

    On the other hand, this is what Haile-Selassie told the Cleveland Plain Dealer:

    "The current hypothesis, which so many people seem to accept, is that they were ancestral descendents [sic, I'm assuming that's a misquote]- that anamensis gave rise to afarensis," Yohannes Haile-Selassie, expedition co-leader and anthropology curator at the Cleveland museum, said in a phone interview from Addis Ababa. "To test that, we need fossils. That's why we think these specimens are really, really important."

    Sinking A. anamensis

    In their current paper, Haile-Selassie and colleagues conclude the following:

    The Woranso-Mille hominids are significant for understanding the evolutionary history of early Australopithecus, particularly due to critical placement within a previously poorly known time period, 3.5 and 3.8 Myr. They are of paramount importance in testing hypotheses about the ancestor–descendant relationship between Au. anamensis and Au. afarensis. The Woranso-Mille hominids shed some light on whether Au. anamensis and Au. afarensis are two distinct species, or parts of a single evolving lineage undergoing morphological change through time. Dentally they are more similar to Au. anamensis from Allia Bay than to Au. afarensis from Laetoli. However, they also share some derived characters with Au. afarensis from Hadar and Laetoli. Based on the currently available evidence, the Woranso-Mille hominids are temporally and morphologically intermediate between the more primitive Au. anamensis from Allia Bay and the slightly derived Au. afarensis sample from Laetoli (Ward et al., 2001; Kimbel et al., 2006). They appear to potentially represent a transitional population within an anagenetically evolving Au. anamensis-Au. afarensis chronospecies (White et al., 2006; Kimbel et al., 2006) providing further support to the well-established hypothesis of ancestor–descendant relationship between the two species. To test this and other alternative hypotheses rigorously, and elucidate the evolutionary history of early Australopithecus, more complete fossil specimens are needed from the critical time period between 3.6 and 3.8 Myr. However, what appears to be evident with the discovery of new fossils spanning the 4- to 3.5-Myr interval is that morphological differences between Au. anamensis and Au. afarensis do not warrant a species level distinction (emphasis added).

    Buh-zaaaaaaam! All your species are belong to us! Kenyanthropus? We won't even dignify it by using the word. A. anamensis? Sunk like the Bismarck.

    The fundamental debate here is semantic. Everyone seems to agree that anagenesis (that is, gradual evolution over time) is a likely hypothesis for this lineage. Where they disagree is how to handle the taxonomy.

    1. Strict cladists want to name species based on the appearance of unique features (that is, phylogenetic species), in which case A. anamensis is a species, A. afarensis is a later species with new characters, and very possibly we need to resurrect Praeanthropus africanus for the Hadar sample, even if it mostly overlaps, since it has a few characters never found earlier and represents a broader sample of postcranial anatomy, which is entirely unknown at earlier sites.

    2. Strict users of a Wiley-like Evolutionary Species concept always place anagenetic lineages into one species. So, the single lineage hypothesis lumps A. afarensis and A. anamensis together. And as Haile-Selassie and crew go on to point out, we might even lump Ardipithecus, if it's the lineal ancestor of the later hominins.

    3. Not-so-stickly people, which is most everybody, pretty much recognize species along with the crowd. A. anamensis has a history now. It's not just early A. afarensis, because, well, lots of people said so. And after all, you can tell the difference between them if you look carefully.

    What's interesting (at least to me) is to read Kimbel and colleagues' 2006 paper, keeping in mind the "following the crowd" scenario 3. In this light, much of that paper is boundary defense. A. anamensis had already elbowed its way into the textbooks, and the paper recognizes the existing taxonomy without attempting any revision. But the demonstration of anagenesis within A. afarensis would be sure to provoke some strict cladists to name some more species -- a species for Hadar, for example. Kimbel and colleagues reiterated that anagenesis within A. afarensis is expected -- it's part of the species' literature, now. So the paper tried to draw two lines in the sand: on the one hand, A. anamensis is real; and on the other hand, no further distinction within A. afarensis is warranted. Taxonomic containment.

    Just a pelvis away...

    But, drawn in this way, both lines in the sand might be washed away by a single discovery. The present pattern of evidence is mostly dental and mandibular. Woranso-Mille may be only one postcranial specimen away from lacking a bunch of derived postcranial characters that are well-evidenced at Hadar.

    After Ardi, I think this is a serious possibility because of the scope of postcranial innovations at Hadar that are not evidenced anywhere earlier. It could be that all the postcranial traits of Lucy and her kin are lineage-typical, going back all the way to Kanapoi (and don't forget the A. anamensis from the Middle Awash). But we don't know this. Given Ardi, it appears that the adaptive package appeared rapidly, after 4.2 but before 3.5 million years ago. It seems to me that there's every chance that A. anamensis, and possibly the Woranso-Mille sample, hadn't built the whole package yet.

    Ah, now I've gotten down to the end of my notes. I think I'm starting to remember why I didn't put them up at the time:

    While the evidence for bipedality in the earlier A. anamensis is not nearly so extensive as that in A. afarensis, nevertheless it is quite compelling, particularly the KNM-KP 29285 tibia. You'd get pretty long odds betting that the Mille pelvic bones looked very different from Lucy's. I have no information about the pelvis at all, certainly no photos, but it would indeed be a surprise for it to be outside the A. afarensis-A. africanus range of variation.

    But then, all it would take is one funky-looking pelvis to throw the whole question of pre-4.0-million-year-old hominids wide open. So maybe we should hope that it's strange.

    Well, we certainly got one funky-looking pelvis, didn't we? I'm beginning to think I should republish old notes more often. What are the chances that another funky pelvis is waiting to be published?

    Could it be that Woranso-Mille could represent an intermediate postcranial form at 3.7 million years? That would be one good reason to nail down the question of anagenesis from the craniodental perspective.

    I think we may already have a hint at the answer. It's a little hard to imagine that Haile-Selassie and colleagues would propose sinking A. anamensis if they already knew that their skeleton has a different postcranial anatomy than represented at Hadar.

    There's one more thing worth mentioning: this paper doesn't include any discussion, comparison -- it doesn't even breathe the name -- of the other 3.5-million-year-old hominin. It's not just a skull; there is a sample of teeth from that unmentioned site, which of course may or may not represent the same taxon. Like a forgotten stepchild of paleoanthropology. Is it possible that peer reviewers have already forgotten it's existence?

    When I wrote about what I'm wondering, well, this isn't the only paper to have recently omitted this obvious comparison. I'll have more on that little problem later on...

    References:

    Haile-Selassie Y, Saylor BZ, Deino A, Alene M, Latimer BM. 2010. New hominid fossils from Woranso-Mille (Central Afar, Ethiopia) and taxonomy of early Australopithecus. Am J Phys Anthropol (in press) doi:10.1002/ajpa.21159

    Kimbel WH, Lockwood CA, Ward CV, Leakey MG, Rak Y, Johanson DC. 2006. Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record. J Hum Evol 51:134-152. doi:10.1016/j.jhevol.2006.02.003

    Synopsis: 
    This may be the snarkiest post about fossil hominins that I've ever hit the "publish" button on.
  • Lomekwi muckraking

    Tue, 2009-11-03 08:38 -- John Hawks

    Michael Balter asks a question I've hit here a few times: "What ever happened to Kenyanthropus platyops?"

    When the talk was thrown open for discussion, [Tim] White took the microphone and began firing questions at Spoor about the degree of variation of the cheekbone position among specimens of A. afarensis and other hominin species. “We took that into account,” Spoor responded, “and I just showed you a graph” about it. “I didn’t ask you whether you took it into account; I asked you what it was,” White said. Spoor, clearly frustrated, told the audience that he had no vested interest in this debate.

    Hmmm...here's an idle thought: Kenyanthropus differs from Australopithecus in having smaller molars (especially first molars), a flat lower face, and small (chimpanzee-sized) ear openings. Ardipithecus differs from Australopithecus in having small molars, big canines, and small ear openings. We now know from Ardipithecus that a hominin-like base of the skull does not reflect obligate bipedality. White argues that the face of KNM-WT 40000 is distorted, so that the "flat face" may be an illusion. There's not a single measurable canine in the Lomekwi (Kenyanthropus) sample, nor are there any postcrania.

    Could KNM-WT 40000 be a late-surviving Ardipithecus? Nah, doesn't work. Dentally, the Lomekwi sample is basically like A. afarensis, except for having smaller molars. Ardipithecus has really small third molars compared to Australopithecus, and lacks the occlusal anatomy of the later hominins. And those small ear openings are shared with Australopithecus anamensis, so they're not probative.

    Still, it reminds us that there may be some shaking-out to do over the phylogeny of these early Pliocene hominins.

  • Is a lack of fossils the problem with early Homo?

    Sat, 2007-09-22 22:55 -- John Hawks

    Just noticing, in this John Noble Wilford article:

    A new report, to be published Thursday in Nature, will review more skeletal evidence of the transitional aspects of the Dmanisi specimens.

    More later...

    UPDATE(2007/09/18): Wilford doesn't directly state the article's theme but it clearly has one: Why the heck can't these people agree about these fossils that have been out of the ground for thirty years?

    The first answer that everyone has given him is about the "million year gap" between 3 million and 2 million years ago. People can't agree about early Homo because they can't decide what its ancestors looked like. Without any ancestors, they don't know which of the traits of early Homo are derived.

    For a good example, we can turn to a feature Wilford doesn't mention: limb proportions. Recently, a lot of ink has been spilled discussing the evolution of arm size in later australopithecines and early Homo. OH 62 (probably Homo habilis) and A. africanus have been argued to have large arms compared to their legs. A. afarensis and Nariokotome (KNM-WT 15000, probably Homo erectus) have relatively small arms compared to their legs. Did H. habilis and H. erectus have different ancestors? Did H. erectus evolve from H. habilis, reverting its limb proportions to earlier A. afarensis? Or are all these comparisons just batty, since only three specimens have arm and leg elements whose length can be compared? There's no clear answer; but one of the most important specimens in the question (with sort-of-intermediate limb proportions) is the Bouri skeleton, BOU-VP 12/1, which at 2.5 million years old is right in the middle of that "gap."

    The more you look at the "gap," the less gap-like it looks. For one thing, we have a pretty good idea of what was going on behaviorally during that million year span. The first stone tools are 2.6 million years old. The technology of these toolmakers -- although simple -- included all the basic manufacturing methods used before 1.5 million years ago. The tools were used to butcher animals and break bones for marrow; so we know that the toolmakers were depending on meat.

    Second, we actually have quite a lot of fossils from this time period. The entire South African A. africanus fossil record, with the exception of a few early specimens like STW 573, come from this "gap." A fairly extensive record of the appearance and evolution of early robust australopithecines comes from this time period in East Africa.

    And, here and there, a few specimens look Homo-like. Wilford's article discusses AL 666-1. To this we can add the Uraha mandible, Omo 75-14, an additional series of teeth from Omo, and possibly the Bouri BOU-VP 35/1 skeleton.

    Properly considered, the rarity of early Homo in these contexts is not a problem; it is information. Wilford quotes Philip Rightmire to this effect, and we can easily expand on the basic concept. Early toolmakers did not undergo an immediate geographic expansion upon their origin. They spread across a relatively narrow strip of East Africa and stayed there for more than a half-million years. They were initially rare. That means that their adaptation was not immediately a barnburner of a success -- the early toolmakers took a while to perfect the adaptation of later Homo.

    The middle part of the article takes in another reason for disagreement: whether H. habilis and H. erectus were ancestor-descendant:

    Several scientists, notably Dr. White of Berkeley, took issue with the interpretation seeming to imply that evidence for the two species overlapping in time and exhibiting variable sizes was new. That, he said, had been recognized for a couple of decades.

    Dr. Kimbel, who was not involved in the new research, defended the authors, saying that they had not "meant to imply that habilis could not have been ancestral to erectus, presumably on the basis of their being contemporaneous at Turkana," the site in Kenya where the fossils were found.

    Susan C. Anton, an anthropologist at New York University who was a member of the Spoor-Leakey team, said, "My money is still on habilis as the potential ancestor, but there is a lot of room for additional knowledge, given the dearth of fossils."

    None of these statements really disagree with each other. If anything, this particular question may have gotten easier to resolve lately, not as a consequence of new fossils, but as a result of new dates for many of the old ones. Susan Anton is later quoted saying that anagenesis "is the only option that is no longer on the table," and it seems to me that this is the clearest statement most likely to invite some hypothesis testing. But it is fairly clear that this problem cannot be resolved in terms of earlier fossils: I don't think there's any compelling evidence of H. erectus before 1.6 million years ago.

    There is one significant word that doesn't appear in the article -- an absence that is especially interesting considering the quoted scientists:

    Kenyanthropus

    Remember, the dominant theme is about complexity and bushiness. And yet, here's that forgotten branch of the family tree; the one that was supposed to clarify everything by providing a different ancestor for KNM-ER 1470 from other H. habilis specimens, the one that showed a distinct line leading to Homo originating in the Early Pliocene.

    I think our bush may have been pruned.

  • Miocene hominids and a crisis of confidence

    Fri, 2007-06-29 12:47 -- John Hawks

    Out of this week's Science Times special on evolution, I clicked into John Noble Wilford's article first, titled "The Human Family Tree Has Become a Bush With Many Branches".

    Now, I don't know about you, but that seems like a boring headline to me. They've been talking about human evolution being a bush for going on 20 years now. It was an old idea when I was in graduate school. So it seems like, if this is all we have going on, the "new frontier" of paleoanthropology must be pretty dull.

    The writer doesn't write the headlines, and the headline doesn't describe Wilford's story, which is basically a verbal slide show of fossil discoveries over the last decade or so. Some bone pictures (of the actual species discussed) accompany the article, and it's a good enough sort of account of new finds since 1990, framed around the tension between fossil finders and molecule mavens.

    But I'll be a little critical. The thesis is that paleoanthropologists suffered a crisis of confidence after molecular data came online in the 1980's, and "a rapid succession of fossil discoveries since the early 1990's has restored" it.

    Well, OK, maybe. But consider the listed discoveries: Kenyanthropus, Ardipithecus ramidus, Ardipithecus ramidus, Orrorin tugenensis, Sahelanthropus tchadensis, Homo floresiensis, and Australopithecus anamensis. Of all of these, only Ar. ramidus and Au. anamensis have gone without significant controversy.

    We can set aside H. floresiensis for a moment -- the controversy about it being possibly the loudest, it also stands apart as the only species listed younger than 3.9 million years. All of these early Pliocene and Miocene species have also been challenged -- by the discoverers of the others, by old hands, and by young upstarts like me. At least one research group has claimed that all of the Miocene "genera" may actually belong to one species. Another has claimed that most of these "hominids" may actually be apes.

    Whether there was any crisis of confidence among paleoanthropologists, all this disagreement is certainly business as usual.

    And, contrary to the article, every one of these species would be thrown from the hominid line, if we believe the molecules. Here's the text from the article:

    Genetic clues also set the approximate time of the divergence of the human lineage from a common ancestor with apes: between six million and eight million years ago.

    Fossil researchers were skeptical at first, a reaction colored perhaps by their dismay at finding scientific poachers on their turf. These paleoanthropologists contended that the biologists' "molecular clocks" were unreliable, and in some cases they were, though apparently not to a significant degree.

    ...

    The new finds have filled in some of the yawning gaps in the fossil record. They have doubled the record's time span from 3.5 million back almost to 7 million years ago and more than doubled the number of earliest known hominid species. The teeth and bone fragments suggest the form -- the morphology -- of these ancestors that lived presumably just this side of the human-ape split.

    It is true that the new fossils date as far back as 7 million years; with Sahelanthropus just under that date, Orrorin at around 6 million, Ar. kadabba at 5.5, Ar. ramidus at 4.4, and Au. anamensis at around 4.1.

    But it has been many years since a genetic comparison indicated a human-chimpanzee common ancestor as old as 6-8 million years. This year's study by Holbolth et al. (2007) estimated a human-chimpanzee speciation time of 4.1 +/- 0.4 million years. That makes Au. anamensis possibly too young to be a hominid. The rest of those species would presumably be just so many apes.

    Now, I don't believe for a second that Au. anamensis is an ape and not a hominid. It just looks too much like Au. afarensis -- so much so that some would put them in the same species. The evolutionary transition between these two is well documented, and will be more so when some as-yet-unpublished fossils come out. So anything younger than 4.1 million years is almost certainly not right for the human-chimpanzee divergence.

    But the 4.1 million year estimate is not unusual compared to other recent studies. My post from last May covers many of these recent studies, including last year's problematic "hominid-chimpanzee hybrid speciation" paper by Nick Patterson and colleagues. The conclusion in that paper about hybridization was certainly wrong, but the date of 5 million years was right in line with other estimates.

    These genetic comparisons are not easily dismissed. Possibly there has been a rate deceleration of mutations in the human lineage that means that the estimated dates are too recent. Maybe 4.1 million years can be stretched into 6 million. Maybe it can even be stretched into 7 million. But all this stretching does have other effects -- on the estimated dates of earlier divergences -- and those are compounded by a large multiple of the few million years we may try to push the human-chimpanzee speciation date. That 4.1 million year estimate is calibrated from an African-Asian great ape divergence at 18 million years ago. Push the human-chimpanzee divergence to 7 million, and you push the orangutan-human divergence back into the Oligocene. Are silent sites in humans evolving more slowly than cercopithecines? Probably. Are they evolving that much slower than orangutans? I suppose nothing is impossible, but maybe we should take another look at those fossils.

    All this is to point out that there really is a conflict between these Miocene "hominids" and genomic evidence about human-chimpanzee speciation time. I don't see any magic solution to this problem from the molecular side -- those dates keep coming up again and again from different regions, and from comparisons across many regions -- including estimates that are not calibrated by other fossil divergences. This is not an easy "the molecular clock must be wrong" kind of problem.

    Nor are the fossils an easy problem. There is pretty good evidence for vertical posture or hindlimb-dominant movement in all of these "hominids." Up to now, we've accepted these kinds of features as de facto evidence of bipedality, and assumed that bipedality is such a unique character of hominids that it is unlikely to be any older. Yet few of these fossils provide really good evidence for obligate bipedality, and some of them provide none at all.

    Is it possible that bipedal apes long preceded the divergence of humans and chimpanzees? Was the common ancestor of the two lineages a biped? Or was significant vertical posture a common feature of many Miocene apes -- making Sahelanthropus a possible homologue of Oreopithecus?

    Which feature is the important one? The long nuchal plane of Sahelanthropus? The femur neck cortical bone distribution of Orrorin? The toe bone of Ar. kadabba? Heck, I can hardly convince my undergraduates about that toe bone!

    I've talked to people about this. Some think that all the molecular stuff is just jibberjabbing, and any day now we will find out that the date estimates were wrong all along.

    I think it may be time to start doubting our confidence again.

    UPDATE (6/28/2007): I've gotten into rather an interesting e-mail discussion about whether I should have included Homo georgicus on the list of new species. Frankly it didn't occur to me: Wilford didn't mention it.

    Actually if you start to think about all the new names that have been proposed in the last 15 years, it is a quite bushy list. It will be no surprise that I think this bushiness has more to do with the listers than the listees.

    Anyway, there is something interesting about early Homo right now that goes beyond the simple splitter/lumper questions. I'll have more to say about it in a few days.

    References:

    Hobolth A, Christensen OF, Mailund T, Schierup MH. 2007. Genomic relationships and speciation times of human, chimpanzee, and gorilla inferred from a coalescent hidden Markov model. PLoS Genet 3:e7. doi:10.1371/journal.pgen.0030007

    Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D. 2006. Genetic evidence for complex speciation of humans and chimpanzees. Nature 441:1103-1108doi:10.1038/nature04789

  • A guide to fantasy science

    Thu, 2007-04-26 22:10 -- John Hawks

    I'm about two-thirds of the way through Mike Morwood's new book, The Discovery of the Hobbit, and I'll be posting a review when I'm through. Generally, I have a positive opinion of the book so far.

    Henry Gee has reviewed the book in this week's issue of Nature. I wanted to point out my generally positive attitude about the book, so that you'll know that my miserable opinion of Gee's review has little to do with the book's merits.

    Consider how Gee starts his review:

    The unicorn, wrote Jorge Luis Borges (in Kafka and His Precursors), is universally regarded as a supernatural being of good omen. But there's a problem: despite its folkloric familiarity, we wouldn't know how to recognize a unicorn if we met one in real life. It "does not figure among the domestic beasts, it is not always easy to find, it does not lend itself to classification," Borges continues. "It is not like the horse or the bull, the wolf or the deer. In such conditions, we could be face to face with a unicorn and not know for certain what it was."

    Is Gee smoking crack? What kind of blather is this?

    First of all, I know I'm being terribly literal, but a unicorn is a horse with a horn. One horn. Not so hard to recognize! Maybe my 3-year-old daughters could help edit at Nature.

    Let's see, where have I seen one of those that Gee might recognize? Oh, yeah:

    UK Pound coins with unicorn prominently visible

    Photo credit: Simon Stratford (via stock.xchng)

    There it is, sound as a pound.

    Next, Gee spends several paragraphs expositing on his own role in the publication of the Homo floresiensis announcement. We learn some interesting little facts, like how the authors wanted to name the species "Sundanthropus floresianus" until a reviewer pointed out that future students would confuse the name with a flowery butt.

    I kid you not. Nature has a layer of reviewers to take tushie references out of taxonomy. Somehow they can't tell a left femur from a right, but they're on the watch for sphincter-species!

    The review is entirely self-serving -- there are only three paragraphs that include any reference to the book! In the midst of this babbling about unicorns and hobbits, Gee tells us that skepticism at new hominid discoveries should be dismissed as the predictable result of "mindsets" of the skeptics:

    Such reaction is common in the wake of new hominid discoveries, which are routinely dismissed either as pathological humans (Homo neanderthalensis) or apes (Australopithecus africanus and Sahelanthropus tchadensis). Such reactions say less about the facts than the mindsets of commentators, who might be unwilling to have their comfortable views of the world so forcibly changed. Confronted with what might be a genuine unicorn, many would prefer to see a pantomime horse with a spike glued to its head.

    Ooooh! Since I'm one who has been notably skeptical of Sahelanthropus and have approached H. floresiensis skeptically, I'm obviously a prime target for this paragraph. It is so comfortable to stay in my view of the world where hominids interbreed with each other. Clearly, a bestiary that includes small-brained island bipeds must shake me out of my comfort zone.

    How could I have been so wrong! When every species ever proposed has faced the same resistance? Sure, Tim White says that Kenyanthropus is a glued-together matrix-filled A. afarensis, but that's just his mindset. Or how about Eoanthropus? Sure, Franz Weidenreich thought that it was just a concoction by "English authors," but couldn't he tell that it was more than just a pantomime skull with an orangutan jaw? Why couldn't I see that these petty minds were just holding back the important work of taxonomy!

    No, no, no. You see, if we approach things skeptically, we won't dare to dream about the unicorns:

    The unicorn remains as it always did, frustratingly elusive. This year, the researchers will return to Liang Bua to see if they can discover more. But stories such as this demand a mythological beast altogether less serene. It is as if the researchers had set out to discover some new form of fossil mouse, only to find that they had grabbed a dragon by the tail instead. And as any devotee of Harry Potter will remind you: Draco dormiens nunquam titillandus.

    The theme of the review is perhaps to be expected from Gee, otherwise known as the author of The Science of Middle-Earth. But I find his mixture of fantasy and science to be especially malaprop in the context of the Flores fossils, since with every fantasy word he detracts from the credibility of the journal's review process!

    Some of you will have seen the episode of The Simpsons, titled "Lisa the Skeptic," where Lisa excavates an "angel" from the ground. Here's part of the synopsis from Wikipedia:

    As Homer attempts to get a motor boat, a new shopping mall in Springfield is being built on an area where a large number of fossils were found. Lisa condemns and protests the building of the mall. Thanks to her protest, it prompts the school to conduct an archaeological dig. When Lisa is digging, it reveals a human skeleton with wings. Springfield's residents are convinced it is an angel, and Homer cashes in by moving the skeleton into the family's garage; however, Lisa is skeptical, believing it may not actually be an angel, and even has Stephen Jay Gould test a sample of the skeleton. The next day, Dr Gould runs to the Simpson house and said the tests came out inconclusive and after Lisa on television compares belief in angels to belief in unicorns and leprechauns, Springfield's religious zealots riot and destroy all of the scientific institutions.

    Later, we find out that the "angel" is a publicity stunt for the new mall; Guest voice Gould confesses that he never really performed any tests on the "angel". This is one of my favorite episodes: it's a rare one where Lisa's preachy skepticism is entirely justified, and the "expert" doesn't care enough to do anything at all.

    Now I know, that the episode was missing a scientific editor to encourage Lisa to forget about her doubts, and just to accept the "angel" for what it is. After all, every new discovery has its skeptics.

    Well, there is a lesson to take away from all the unicorn talk. If you are in Cardiff and find the skeleton of a giant, be sure to send your report to Nature, where you'll find a receptive editor. Despite what they may say, there's not one of those born every minute.

    UPDATE (4/26/2007): A reader e-mails, "Remember that Borges was blind." True. Perhaps we can extend this analogy further?

    Another reader: "Well, at least we can expect a fair set of reviews on the Sahelanthropus postcrania...D'oh!"

    References:

    Gee H. 2007. In a hole in the ground.... Nature 446:979-980. doi:10.1038/446979a

  • Commentaries on Dikika

    Fri, 2006-09-29 14:44 -- John Hawks

    The online companion site to Scientific American is running commentaries by a few paleoanthropologists on the importance of the new DIK-1-1 skeleton. It's an experiment in online publishing, and it has turned out some very good material, including commentaries by Owen Lovejoy, Ralph Holloway, Diana Roman, and, well, me.

    These other commentators and the press of the last week are hard to follow, but actually my second thought about the skeleton has turned out to be fairly original:

    The new Dikika skeleton has me wondering one thing: did Selam sink Kenyanthropus?

    If you can guess the connection, you've been reading entirely too much about early hominids!

    Congratulations to Kate Wong on a really innovative idea -- they have an opportunity for other folks to contribute as well, so dive in!

    Synopsis: 
    Along with others, I contribute a perspective on the Dikika child in Scientific American.
  • Our quondam homs

    Wed, 2006-09-20 23:25 -- John Hawks

    Did I miss a meeting?

    Thanks to efforts in Ethiopia and elsewhere, we already know a good deal about A. afarensis. It has been called an 'archaic' hominin for at least two reasons. First, it is old: its fossils date from between 4 million and 3 million years ago. Second, its morphology is archaic, in the sense that its brain case, jaws and limb bones are much more ape-like than those of later taxa that are rightly included in our own genus, Homo. When adjusted for its body size, the brain of A. afarensis is not much larger than that of a chimpanzee, and although it has lost the large canines that distinguish apes from hominins, other aspects of its dentition, such as its relatively large chewing teeth, are still primitive (Fig. 1) (Wood 2006:278).

    Every other reference on the internet to "archaic" hominins, hominids, or homininos refers to Middle Pleistocene Homo. So what's going on with this?

    I guess that "australopithecine" no longer appeals to folks who want to simultaneously refer to Australopithecus, Kenyanthropus, Ardipithecus, Orrorin, Paranthropus, and whoknowswhatelseensis. So maybe some people are casting around for another term, besides the boring "early hominid" -- oops, "hominin".

    It doesn't make sense to redefine "archaic" to mean non-Homo hominids -- oops, hominins. So I thought I would look in my thesaurus for some alternatives:

    age-old, aged, antediluvian, antiquated, antique, archaic, back number*, been around*, bygone, creak, dated, decayed, done, démodé, early, elderly, erstwhile, fossil*, hoary, moth-eaten*, obsolete, old goat*, old-fashioned, older, oldie*, out-of-date, outmoded, primal, primeval, primordial, quondom, relic, remote, rusty, sometime, stale, superannuated, timeworn, unfashionable, venerable, vintage

    Now, sure "fossil" is out -- but there are a lot of good options here. I think "hoary hominids" is a bit catchier than "old goat hominids", er, "hominins". But maybe "quondam hominins" is the way to go.

    References:

    Wood B. 2006. A precious little bundle. Nature 443:278-281. Full text (free)

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