john hawks weblog

paleoanthropology, genetics and evolution

Hadar

  • A Lucy remembrance

    Sat, 2011-09-10 13:12 -- John Hawks

    The CNN medical blog (associated with Sanjay Gupta) is running a short piece by Don Johanson, which may be of interest: "'Lucy' discoverer: Why I study human evolution".

    My deep commitment to understand the origins of humankind was ignited when I read Thomas Henry Huxley’s 1863 book "Man’s Place in Nature." The core idea that gripped my teenage mind was the suggestion that humans and African apes shared a common ancestor that roamed Africa millions of years ago.

    I think if you're a teenager reading Huxley, you're already in the right tail of the distribution for "human evolution interest"!

  • Through the early Homo archives

    Mon, 2011-08-29 22:32 -- John Hawks

    I've enabled the search function for the site, which you'll find at top right on each page of the site. The search index is still rebuilding, and as I write this has only indexed 4% of the site. That brings it up to late 2007, and it's interesting to go back through the history of paleoanthropology that way.

    For example, I ran across my comments ("Is a lack of fossils the problem with early Homo?") on a John Noble Wilford piece from four years ago. Seems very timely in many ways. For example, the paucity of the fossil record of Homo before 1.6 million years ago was a major feature of the article. I directed my attention to the supposed "gap" between 3 and 2 million years ago:

    [W]e 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.

    Of course, dates have changed. We now have good dates for Dmanisi, which make those fossils the earliest well-attested Homo erectus sample at 1.8 million years. STW 573 now looks late, not early. But the fact remains that people were looking for pure representatives of Homo in the fragments instead of exploring morphological diversity within the large and fairly complete samples at hand.

  • 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.
  • Lucy Seattle review

    Fri, 2009-03-13 07:51 -- John Hawks

    The NY Times has a review of the Pacific Science Center's Lucy experience, which came to an end this week. They're blaming the financial loss on Obama:

    But the sour economy does not seem to explain all of Lucy’s troubles. A rare December snowstorm played a role, and Bryce Seidl, the center’s president and chief executive, has suggested less intuitive reasons like the feverish focus this liberal city had on the election of President Obama and his transition to office.

    Yep. That's gotta be it. Meanwhile,

    “This is going to be a big thing,” [Ethiopian honorary consul] Mr. Kebede said. Next year, he said, he hopes Lucy will travel to China for an exposition in Shanghai.

  • Mysteries of A. afarensis

    Mon, 2008-10-06 12:00 -- John Hawks

    In honor of Lucy's move to Seattle, Alan Boyle has a piece at "Cosmic Log" about Lucy and A. afarensis. It has a lot of questions and few answers, but may be interesting for students. And there's this passage involving Anna Behrensmeyer:

    Even Johanson has said the First Family is as deserving of fame as Lucy. According to South African reports, he's trying to nail down the evidence for the cause of death by consulting with an expert on prehistoric predators.

    In the past, investigators have suggested that the entire troop of hominids might have died in a flash flood, or were done in by a bout of food poisoning. But Behrensmeyer said the current prevailing theory is that they were the victims of a "surplus killing" by blood-crazed predators. "This is documented as a natural phenomenon," she said.

  • Paleoecology at Hadar

    Thu, 2008-03-20 12:23 -- John Hawks

    The coming attractions bin at Journal of Human Evolution includes a paper by Kaye Reed, reviewing the evidence of paleoenvironment in the Hadar formation:

    Habitat reconstructions of 12 submembers of the Hadar and Busidima formations (˜3.8-2.35 Ma) are presented here along with faunal differences in these submembers through time. Habitats with medium density tree and bush cover dominated the landscape through much of the earlier time period in the Hadar Formation. The lowermost Sidi Hakoma Member is the most closed habitat. The Denen Dora Member shows the influence of frequent floodplain edaphic grasslands with high abundances of reducin bovids. There is an influx of ungulates in the Kada Hadar Member (˜3.2-˜2.96 Ma) that indicates a more arid habitat populated by mammals that were recovered from earlier deposits further south in Ethiopia and Kenya. In the younger deposits from the Busidima Formation at Hadar, the landscape was open wooded grassland with some floodplain environments. The fossil assemblages from the Busidima Formation show a substantial species turnover. Although high numbers of A. afarensis specimens are associated with the lower Sidi Hakoma Member, they clearly inhabited a variety of habitats throughout the entire Hadar Formation. Australopithecus afarensis from Laetoli through Hadar times appears to have been a eurytopic species.

    This is a nicely detailed paper, focusing on the amount of wooded/bush habitat, the relation of the hominids to those habitats, and the relative lack of early faunal exchanges with areas further to the south.

    The discussion focuses on the range of paleoecologies in which fossil A. afarensis has been found -- including not only Hadar but also nearby Maka and Dikika, and more distant Koobi Fora and Laetoli. Altogether, these localities cover a long time (from before 3.5 up to around 2.9 million years ago). From the range of paleoecologies reconstructed in this paper at Hadar, Reed concludes that A. afarensis did not have a "narrow" habitat preference. It is found in relatively closed woodland, open woodland/bush, and wet grassland/marshland.

    There are some differences between localities. At Koobi Fora, relatively few specimens of A. afarensis have been found in the Tulu Bor Member, despite the fact that it occupies the same time as the Hadar sequence. Based on the paleoecological data, Reed suggests that Hadar was a wetter, more closed woodland habitat than Koobi Fora at that time -- Koobi Fora would have included more scrubland punctuated with wetlands and floodplains (here she cites her own 1997 paper).

    The early end of the A. afarensis sample is represented at Laetoli. Reed gives a brief review of the paleoecology of that site, which has been interpreted differently by different authors but broadly appears to have had a fairly high amount of rainfall and some patches of forest amid closed woodland:

    Thus, the earliest known A. afarensis material was found in deposits showing habitats in which trees and or bushes were fairly plentiful. It is also interesting to note that while the deposits of A. afarensis at Laetoli and Hadar share some perissodactyls, giraffids, suids, and proboscideans, the bovid taxa and those primates other than A. afarensis are not very similar.

    Reed concludes that A. afarensis was a "eurytopic" species -- one that inhabited a wide range of habitats and moved broadly across space. It contrasts with the more habitat-selective ("stenotopic") species, which include most of the bovids.

    White et al. (1993) suggested broad habitat tolerance for A. afarensis, and indeed, the species has thus far been recovered from regions in which the reconstructed habitat ranges from closed woodland through more open, but wet woodland and shrubland. There is no direct evidence that A. afarensis only existed in riverine forests or grassland habitats, or that they preferred one habitat over another. It is tempting to equate the aridification in the Kada Hadar Member with the extinction of A. afarensis. However, sediments at Hadar are sparse or missing altogether from ˜2.90-2.35 Ma thus obscuring details of the species' demise. All that can be said is that they are no longer present at 2.35 Ma and most of the fauna, including hominins, has been replaced.

    References:

    Reed KE. 2008. Paleoecological patterns at the Hadar hominin site, Afar Regional State, Ethiopia. J Hum Evol (in press) doi:10.1016/j.jhevol.2007.08.013

  • Building a research infrastructure in Ethiopia

    Sat, 2008-03-01 17:27 -- John Hawks

    In the current Science, Elizabeth Pennisi reports on Ethiopia's efforts to bring more resources and tourism to its fossil heritage:

    Then in 2003, that lab [at the National Museum, built in 1982] was razed to make way for a six-story, modern structure that includes a two-floor library, a 500-person auditorium, and 200 rent-free offices, plus storage and study space for more than a million specimens. The three wings are devoted to paleontology and archaeology; art and history; and administrative, conservation, and educational spaces.

    ...

    Foreign aid is helping: France is supplying furniture, and Japan may outfit the hominid spaces. Everyone involved is thrilled and not just with the prospect of more space. "It shows how much emphasis has been given [to research]," says Ethiopian native and paleoanthropologist Yohannes Haile-Selassie of the Cleveland Museum of Natural History in Ohio. "In a country that has a lot of needs, the government could have easily used that money for something else."

    The article includes a map showing the impressive array of 25 active archaeological or paleontological field sites across the country. The funding and effort are also being devoted to training students -- not only in paleontology but across the sciences -- including the establishment of new universities.

    One of the more controversial areas is tourism -- including the encouragement of travel to active research areas:

    At the same time, "one of the most important things that needs to happen is the integration of tourism and science," says [anthropologist Tim] White. And that, too, is happening. National Geographic has pledged support for an educational center at the village nearest to Hadar, home of the 3.2-million-year-old Lucy. With better roads under construction, "it could easily be a destination spot for tourists," [anthropologist Donald] Johanson predicts. Exhibit plans are still taking shape, but there likely will be casts of Lucy and other fossil hominids, as well as photographs from the site.

    This raises fears (by some) that the sites will be more vulnerable to destruction by trampling and looting. On the other hand, bringing a better transportation infrastructure is bound to improve matters for the scientific research teams, possibly including the most enduring problem -- security.

    Personally, I wonder whether such projects can be a financial success. Many of the paleontological museums in the American West are a great experience for visitors because you can get close to the original (and often still-active) sites, see original specimens, and do it all without the huge crowds of urban natural history museums. But that's the point: there are no huge crowds of people. These are substantial tourist draws for small towns in the West, but they aren't often making back the substantial federal or state grants that help to build them. It's an important cultural resource and a valuable investment, but it may not be realistic to expect small regional Ethiopian museums to draw premium paleo-tourist dollars.

    References:

    Pennisi E. 2008. Rocking the cradle of humanity. Science 319:1182-1183. doi:10.1126/science.319.5867.1182

  • An interview with Michelle Drapeau

    Sat, 2008-01-12 20:17 -- John Hawks

    I've been trying to spread the interviews across the field in various directions. I (virtually) talked with Mica Glantz about Neandertals, Adam Van Arsdale about early Homo, and Anne Weaver about human brain evolution, all the australopithephiles in the readership are probably feeling neglected.

    So I wrote to Michelle Drapeau, who was very generous in answering questions about her work on the anatomy of early hominids and her recent field work in Ethiopia. Michelle is on the faculty of the Université de Montréal, in the Department of Anthropology. She serves as co-director of field operations in the Bala Paleoanthropological Research Area of southern Ethiopia.

    Hawks: I will start out by asking about your dissertation work, which centered on the new partial skeleton from Hadar, A.L. 438-1. How did you get involved in that analysis?

    Drapeau: It's a case of being at the right place at the right time. Bill Kimbel and Don Johanson had asked my advisor at the time, Carol Ward, to describe all the postcranial material recovered from the field in Hadar since 1990. Among those specimens was the partial skeleton of A.L. 438-1 which included associated fragments of the humerus, clavicle, radius, right ulna, mandible, and frontal as well as a complete left ulna, right and left second metacarpals and left third metacarpal. Considering the relatively numerous body parts from one individual, Carol thought the specimen deserved a more detailed analysis. I was Carol's Ph.D. student at the time and the 438-skeleton (as we started to call it) appeared like an ideal subject.

    Hawks: What did you have to learn to be able to undertake the work?

    Drapeau: I had to learn a lot! My master's thesis was in the history of science field, so all the functional anatomy, including the descriptive and comparative aspects were completely new to me. It was something I really wanted to do, however, so I really enjoyed immersing myself into it.

    Hawks: A.L. 438-1 exhibits more curvature across its length than A.L. 288-1, an issue that you discussed in your analysis of the fossil. I have always been puzzled by the problem of ulna curvature -- mainly because I've always been puzzled by the comparison of later, larger, and more curved fossils like Omo L40-19 and OH 36 -- and then, of course, KNM-WT 15000 is a lot more like most recent humans. Do you have any insights about these contrasting morphologies?

    Drapeau: Forearm bone curvature is an intriguing issue. Intuitively, it makes sense to assume that curvature reflects arboreality since the curvature of both the ulna and radius give greater area on the interosseous membrane for attachment of forearm muscle important for arboreal locomotion such as the finger flexors. However, orangutans and gibbons do not have the most curved forearm bones. It is an honor that goes to gorillas, definitely not the most arboreal animal of the bunch. If the area of muscle attachment is the variable that interests us, then it is important to take into account forearm length as well. When that is done, species generally sort by locomotor preference, with the most arboreal having the greater ‘area' for muscle attachment relative to body size and humans having the smallest (at least, when measured on the ulna). So gorillas appear to have very curved forearm bones because they also have relatively short forearms when compared to other apes.

    The differences between A.L. 438-1 and A.L. 288-1 are fairly minor and probably reflect normal within-species variation. Neither is very curved and they may belong to a population with slightly more curved ulnae than modern humans but definitely less curved than any extant apes.

    The KNM-WT 15000 specimen is pretty much what you would expect an ulna belonging to a completely terrestrial biped to look like, i.e., it is not particularly curved. Since it is a juvenile, it is difficult to compare it to other fossils, but there is nothing really surprising about it.

    That said, what about the intriguing Omo L40-19 and OH 36? These specimens present combination of morphologies that are difficult to underscore in quantitative analyses. The former had a human-like proximal morphology but a really long and curved (ape-like) diaphysis. The latter, OH 36, has a general ape-like morphology with a pronounced curvature, but is unique in a few characters. The whole bone (proximal articulation and diaphysis) is very constricted medio-laterally, more comparable what is observed in monkeys (and it is not the result of distorsion). Despite its general ape-like morphology, it has an olecranon process that projects proximally like no other ape of its size. It is definitely much more human-like for that trait and it is generally agreed that it is a hominin. McHenry and colleagues argue in a recent article (AJPA, 134: 209-218) that these two fossils are very different and can hardly be accommodated into the same genus (Paranthropus) as it is usually done (probably by default). McHenry and colleagues argue that it may indicate Paranthropus is in fact a polyphyletic taxon. They also conclude, as I stated above, that OH 36 is unlike anything living today.

    So, if curvature of the ulna reflects arboreality, does it mean that these fairly recent fossils were much more arboreal than A. afarensis? Remember that they are big ulnae, particularly L40-19, likely belonging to large individuals.... Maybe the Paranthropus clade (if indeed it is a clade) is more arboreal than A. afarensis? This would imply either reversal of behavior or that A. afarensis is not ancestral to Paranthropus. Or, alternatively, could the curvature in these individuals reflect forelimb muscularity but not necessarily related to arboreality? As you can see, I have many more questions than answers. All this variability suggests that the behaviors of fossil hominin species were much more variable than what we have been used to think and may have been (very?) different from the behaviors of extent species.

    Hawks: Of course, the big debate about forelimb proportions is the idea that they may have been very different (and more apelike) in A. africanus compared to A. afarensis. (reviewed by Green, Gordon, and Richmond 2007) What do you think about the issue?

    Drapeau: That idea first met with some resistance because it involved a reversal of proportions from A. afarensis to A. africanus and implied a more arboreal behavior in the latter than the former. Given that Homo habilis is often described has having more ape-like proportions than A. afarensis, it also implied that A. afarensis may not be the ancestor of the Homo lineage (an idea more recently suggested by Yoel Rak and colleagues based on mandibular data). Since I remain unconvinced of the primitive proportion of H. habilis, I am not so certain that the 'derived' proportions of A. afarensis exclude it from being an ancestor to the Homo lineage.

    Back to the differences between the two australopithecine species. Despite original skepticism, the data appears to be robust and the differences in joint size between A. afarensis and A. africanus appear to be real. As observed in the previous question, this variability may reflect locomotor differences possibly related to differences in the environment. If A. afarensis was still occasionally arboreal, is it too hard to imagine that, if the environment is changed (more wooded, greater predator pressure, more resources found in trees, etc.), the percentage of arboreal behavior would increase and that the proportions would revert to being more chimp-like in A. africanus? Again, there is no reason to assume that all early hominins, because they were bipedal, were identical in their locomotor behaviors.

    I want to underscore that these differences are in joint SIZE, not in limb length, and reflect relative loading of the limbs. Usually, the major source of loading of the limbs is related to locomotion, but it is an assumption that cannot be verified in early hominins. If, as stated above, OH 36 is unlike anything living today, maybe it did things that have no modern equivalent. And the same can be said of other hominin species including A. africanus with its 'apparent' primitive proportions.

    Hawks: You have recently been involved in field research in the Bala-Weyto region of southern Ethiopia. Can you describe the site, and your role?

    Drapeau: The Bala–Weyto basin is part of a series of small parallel rifts that link the northern limit of the East African Rift to the southern limit of the Main Ethiopian rift. These small rifts constitute today a string of many small basins. The Bala-Weyto basin is located east of the Omo river basin. It is a region more difficult to survey when compared to dryer region because of the vegetation coverage that limits exposures visibility and access. However, it is little-explored paleoanthropologically speaking. Work in the Konso, another small basin a few kilometers away, but at a higher altitude, has a fauna with a certain degree of endemism and an A. (P.) boisei specimen with unique morphological variations. Among other things, we want to know if this variation and the faunal endemism are due to the relative isolation of the basin or to its particular environment. These answers may be found in contiguous basins that vary in their physical characteristics, such as the Bala-Weyto basin.

    I am co-director of that project with Elizabeth Harmon of the City University of New York. At this stage of the project, being co-director involves organizing the whole expedition, securing funding, and coordinating the work of other team members. I would say that the most time consuming aspect is coming up with money and getting everything moving in the field. As a director, I am responsible for the team's well-being and it is a pressure that can sometimes weigh heavily on my shoulders. It is nice to be able to share the burden with a co-director.

    Hawks: Do you involve students in your work?

    Drapeau: My funding is limited and field work in Ethiopia is not particularly cheap. However, I plan to bring one student in the field this summer. I look forward to share this experience with a highly motivated student!

    Hawks: Many of us have heard about the difficulties of field research, particularly in East Africa. What are some of your biggest challenges?

    Drapeau: Doing field work in Ethiopia can be a challenge for many reasons. As can be expected, there are numerous permissions, letters, official documents, etc., that are required and the bureaucracy is somewhat heavy. However, I find Ethiopians very helpful and professional and, usually, the quest for documents goes smoothly, particularly once you know what to do and in what order.

    A second difficulty is the access to the sites. Ethiopia did not have one highway until relatively recently and road traveling remains an experience that can be frightening. A lot of work is being done on the roads, however, and I believe that things will keep improving. Access to the research area involves off-road traveling as well, with all the difficulties that it entails. When you leave for the field, you have to be a self-sufficient unit, relying on the local environment as little as possible. It is still necessary to get gasoline on a regular basis, but except that, we try to be as autonomous as possible. It is particularly important when you go to a new area and don't know what (if anything) will be available to you.

    A third aspect of field work, particularly in Ethiopia, is the politics, the paleoanthropological politics that is. Although most scientists are polite and civilized to each other, I really feel that we had to walk on eggs when we were researching an area in which to conduct field work.

    A final difficulty (and certainly not the least) in our situation, is to find an area that has fossiliferous exposures of a time period that interests us and in which we can work at least a few years. The numerous discoveries that are made in East Africa give the impression that finding hominin fossils is something easy to do, but it usually involves many years of surveying. We are still at the exploratory phase of our project, i.e., we are still actively looking for an area that could sustain scientific work for a few years. Hard work (and perhaps a little luck) is essential.

    Hawks: You had a lot of field experience before going to Ethiopia. How did you get your start?

    Drapeau: At the end of my undergraduate degree, I had the chance of getting a couple of paying jobs in prehistoric archaeology. It was the beginning of a series of jobs in field archaeology conducted in parallel to my studies. I used to think (and still do) that these were the best summer jobs an anthropology student could have. The pay check was very descent and it usually came with room and board. These jobs allowed me to see many regions of Quebec and Canada that I would otherwise have never visited and to do things I would probably have never done otherwise. I have flown in helicopters for hours (and even survived a major crash), piloted a hydroplane (just for a few minutes, but still!), hear wolves howl into the night while trying to sleep in a tent hundreds of miles from any road or civilization, dipped my foot in the arctic ocean (too chicken to swim), seen the midnight sun, and I could go on. This fieldwork experience, and a stint in the Caune de l'Arago in Tautavel, France, opened another door: to be invited to do field work in Hadar in 2000.

    Hawks: Any interesting stories?

    Drapeau: I have an anecdote that I find amusing, but mostly informative on the nature of humans. When we were doing field work in the Bala basin, our camp was set up about a 2-hour drive off the road. It was clear that the local people had seen very few foreign workers. For the whole time we were there, we had a constant group of people just sitting in the shade observing us like zoo animals, watching our every move, laughing when we did things unexpected, etc. We were quite the entertainment. The occupation of the local Mali people appeared to be tending their few sorghum fields, but mostly to take their sometime large herds of cows, goats and sheep a few miles down to the river for a drink every day. Even though it was not that hot, the men walk around wearing only colorful underwear (the Speedo-type) and it was sometimes literally falling apart. From our western perspective, they really seem to have almost nothing. Anyhow, after a few days in the field, some crew members were starting to crave fresh meat. We agreed to allow the cook to purchase one goat from a local herder. We didn't think it would be a problem given the large quantities of these animals around and our willingness to pay a fair price for it. It came as quite a surprise that no one was willing to sell us any! It turned out that goats, sheep and cows were not herded to be eaten or even milked, but were really just status items. One man from the village nearby apparently owned more than a hundred head of livestock but was still unwilling to sell. We were all quite shocked of the apparent frivolity of it all, particularly considering that food (for humans and beasts) did not appear to be particularly abundant in the region. But then, we couldn't miss seeing the connection to what we can observe in the western world: huge houses for one or two people, oversized and overpriced cars. These are just to show off. The same frivolities, although expressed slightly differently, can be found anywhere. I guess it really is in the human nature. We were finally able to convince someone to sell us a goat, but we paid a really high price.

    Hawks: Congratulations! You seem to be a very busy person right now, both professionally and personally. What's next for you?

    Drapeau: I just started one of the most challenging projects of my life, a project that will keep me busy for the rest of my life. His name is Henri and he is almost 8 months old. Professionally speaking, I am investigating manipulatory adaptations in the early hominin hands and the morphology of muscle markings. However, one of my main objectives in the next two years is to settle on a specific field research area with good scientific potential.

  • Book review of "The First Human"

    Thu, 2006-06-29 21:27 -- John Hawks

    Despite all the trouble I had traveling (or maybe because of it), I got to have a really enjoyable time finishing Ann Gibbons' new book, The First Human. For a while I was really afraid I'd lost it in the backpack without knowing how it ends! But what a relief, it was in another suitcase so I can report on the whole thing.

    I've read most -- not all -- of the recent trade books about paleoanthropology, and this is definitely one of the top few in terms of being fun to read. It follows a familiar form: the quest for the source of the Nile. The book even mentions Burton and Livingston, whose explorations were to some of the earliest anthropologists what the Leakeys discoveries were to the current generation. Like the quest for the solo transatlantic flight, the summit of Everest, or the race to the Moon, the paleoanthropologists here all are trying to capture the same prize: the earliest hominid.

    The book appeals in large part because it is well-written. Instead of beginning with the long dry history of finding bones in old dry places, Gibbons' first chapter plunges us right into the middle of three discoveries of the mid-1990's -- all happened within six months of each other, but the events of January 1995 brought them together. The chapter even ends with a cliffhanger!

    Then comes the long dry history, with the usual cast of characters: Haeckel, Dubois, Dart, Louis and Mary Leakey. I was apprehensive about this -- no book ever seems to skip this stuff, and it's usually the same boring slog -- but Gibbons adds some details that most people haven't seen before. She's mercifully light on the "Dart courageously fighting the scientific establishment" theme, and brings us a great description of Dart excitedly opening the crate containing the Taung fossils at a friend's wedding. We get rather less of Louis Leakey's long struggle for recognition and more of his behind-the-scenes support from LeGros Clark.

    Most notably, Gibbons brings us sketches of many of the paleontologists that the usual accounts miss. We see Bryan Patterson find not one, but two of the earliest hominids, and the episode that caused him to leave Kenyan field work, with his site of Kanapoi lying fallow for 30 years. We are led down the blind alley of Ramapithecus with Elwyn Simons and David Pilbeam. And we follow Yves Coppens to the Omo, Hadar, and Chad. Indeed, one of the real highlights is the account of field research in Chad, which I haven't seen described elsewhere in English so well.

    The soap opera really begins with the origins and education of the current fieldworkers, who are as interlinked as characters on Days of our Lives. Pilbeam plays a Kevin Bacon-like role connecting Michel Brunet, Andrew Hill, and Martin Pickford. Pickford and Richard Leakey were old schoolmates, and -- maybe or maybe not, according to the book -- Hill comes between them. The chief fossil hunter from Hill's team goes to work for Pickford. The son of the chief fossil hunter for Richard and Maeve Leakey goes to work for Hill.

    We see quite a bit less of the soap opera in Ethiopia, which describes the current Middle Awash work extensively but has little to say about Hadar or other current field sites. Donald Johanson's perspective on events of the last twenty years is very noticeably absent. We see Mary Leakey's anger at White and Johanson for naming her Laetoli discoveries Australopithecus afarensis, but the section does not explain the justification for the anger -- attaching the name to LH 4 as the type specimen removed any chance of naming the Laetoli hominids anything else.

    Ian Tattersall raised an important point in his Nature review of the book: Any reporter who depends on access to subjects faces a possible conflict of interest. Report bad things about the subjects, and they may restrict access. Gibbons has obviously received exceptional access to some of the book's subjects -- indeed, the book mentions the famous lack of journalistic access to some of the research teams. Has this exceptional access affected the narrative?

    I think that the book has a fair account of many events, but omits other well-known incidents that might have been described. For most of these, there is little that Gibbons could have done -- after all, if some subjects don't talk to you, and others won't give details about certain events, then what are you going to write about? In fact, there must be an intense incentive for many people not to cooperate with a book like this, especially those hoping to continue fieldwork in Ethiopia or begin there in the future. The accounts that are in the book make quite plain that one misplaced word can result in field permits being revoked, or access to collections being revoked, or even worse. As a result, the book puts on the record many arguments that were aired in public -- like the dispute over the Galili field site, for instance -- but doesn't necessarily give the whole story.

    There is pretty obviously one overarching prize that shapes the entire narrative. The introductory chapter ends with the world on Alan Walker's "tenterhooks" -- in 1995! -- waiting to see the Ardipithecus skeleton. The book describes on four occasions just how fragile the skeleton was. Twice we hear how the condition of the skeleton "tempered" the Middle Awash team's excitement, twice it is described as "the most fragile skeleton ever found," twice as "roadkill." Early in the book White emerges as a secretive Svengali; at the end -- during an event White himself describes as "theater" -- we see him casting aside the velvet curtains to show his specimen at last to his skeptical colleagues.

    Except, well, we don't get to see it. A reader might be forgiven for thinking the obviously crushed skull on the book jacket is the centerpiece of the book -- its "crushed" skull is twice mentioned. Sadly, no, the cover shot is just Sahelanthropus. Ardipithecus is still locked in its fortress of solitude, unseen by the unwashed. This does raise some concern for me -- since Gibbons will undoubtedly be writing the story of this fossil when it at last surfaces.

    But some of the best moments are those that shine light on the relationship of the science to journals and the media. Two of the major research teams make a point of rejecting the taint of National Geographic and its film crews. In counterpoint, the book repeatedly notes the long association between National Geographic and the Leakey family, including a direct contrast between the histories of Richard and Maeve Leakey and Tim White. Amid descriptions of media-savvy scientists, we see Henry Gee, editor of Nature, commenting on fossils, prognosticating on future discoveries, "prodding" researchers, and having one incredible meeting that was hard for me to believe even after reading it. If one wonders about possible conflicts of interest for Gibbons, how much more must one wonder about the chance of one of these papers being rejected by Nature's vaunted six "peer reviews"?

    At its bottom line, the book really raises two substantive issues. The first is the real danger of today's field work. Paleoanthropology is not merely a game today, it is "the Great Game" replayed. Field teams divide up "Connecticut-sized" research territories, hem opponents into areas with younger sediments, and -- when bullying, scientific name-calling, and bureaucratic manouvers fail -- finally agitate local people, enlist bandits, or pull their guns. To me, the book's most touching moment is its description of Michel Brunet's feelings after losing a colleague on his field team. In another episode, a young graduate student (who deserves recognition for her science and not this) personifies a near-miss with violence in the field. The two cases together bear rereading: if paleoanthropology continues along its current path, then who can doubt that some people will be killed in the field?

    The other issue is the relationship between these field teams and the science as a whole. As depicted in the book, they clearly do resemble explorers looking for the source of the Nile. They know what the goal is -- at one point, Pilbeam even sketches what the ancestor will look like, at another Henry Gee opines about it. It is still out there waiting to be found, and these teams will be searching until they find it. It's "the First Human" of the title.

    But these fossils aren't human -- and it's darned hard to tell whether they are even the more humanlike kind of apes! In the book, we see that the science turns against the scientists sometimes. Ramapithecus is no longer considered hominid by anybody -- it's not even a valid taxon anymore. Louis Leakey's Kenyapithecus wasn't a hominid either.

    Can it be that all of these new fossils are really hominids? Or have some of these scientists in their quest for older and older fossils overshot the mark? The current scientific debate over specimens is only glossed here -- the book sketches what the disagreements are, but gives no details to judge the arguments. (If you want those details, you'll need to read the blog!) Instead, the science appears as another forum for the scientists to misbehave -- accusing each other of holding "creationist positions" and the like.

    Many readers will surely be puzzled to read how these men and women, who brave disease, bullets, broken families and years of denial, can be so poorly composed in the face of scientific examination. Again and again we see them squirrel the fossils away, withdraw them from the world, or give up on paleoanthropology altogether. How can it be that this story is repeated so many times? But the reader should consider: No one can take away Hillary and Norgay's summit photos. But even after all the years of work, the lowliest graduate student might turn one of these "hominids" into an ape.

    Even I make a brief appearance in this book -- blink and you'll miss me dancing through to aggravate Brunet's heart condition.

  • "The catechism about genetic drift"

    Tue, 2006-05-02 13:32 -- John Hawks

    Razib has been working over genetic drift real good (concerning effective population size and population history, and founder effects). It deserves it.

    This post is about genetic drift applied to phenotypic -- not molecular -- evolution. The two are distinct for two important reasons: first, phenotypes are widely genetically correlated with each other while unlinked DNA sequences are not; and second, because the theoretical reasons for some nucleotides to have no correlation with fitness are very strong, but such theoretical reasons are nonexistent for phenotypes.

    Personally, I think selection is more important than drift at the molecular level as well, for reasons having to do with those "genetically correlated" and "unlinked" assumptions. But to the extent that neutral evolution may be credible for many genes, it is much less credible for most phenotypes.

    Here's what I tell my students:

    To explain the evolution of a feature in ancient humans, genetic drift is my absolute last resort...right before sexual selection.

    Why I don't like sexual selection is a topic for another day.

    For now, on to genetic drift. Here's what Gould and Lewontin's famous "spandrels" paper has to say:

    Have we not all heard the catechism about genetic drift: it can only be important in populations so small they are likely to become extinct before playing any sustained evolutionary role (but see Lande 1976) (Gould and Lewontin 1979:585-586).

    With some math, we can show that the "catechism" is not literally true -- genetic drift can cause substantial phenotypic evolution in large populations. But ...

    Some worked examples

    Lande (1976) shows genetic drift can cause phenotypic evolution consistent with many examples in the fossil record:

    This paper presents a statistical test for the hypothesis of evolution by random genetic drift, contingent on the effective population size. In examples from the fossil record, it is found that the rates of evolution equal to or greater than those observed have a significant probability of occurring by random genetic drift even in very large populations (Lande 1976:314-315).

    Let's consider an example not examined by Lande. One of the most significant temporal trends in the early hominid species Australopithecus afarensis is a decrease in the length of the lower third premolar. This decrease in length is associated with a change in the morphology of the tooth, in which a more sectorial one-cusped form becomes less common and a more bicuspid form becomes more common. Lockwood et al. (2000) show that the P3 length decreases from an average around 10.5 mm in the early, 3.5 Ma Laetoli sample down to an average around 8.5 mm in the latest 3.0 Ma Hadar sample. Estimating the standard deviation of the sample as a whole (including intermediate time periods at Hadar) is a bit complicated, but if we consider the mean as a moving average, then the standard deviation is between 1.0 and 1.5 mm. I'll assume 1.0 mm to be conservative.

    Lande (1976) derives the distribution of phenotypic change due to genetic drift in a population with effective size Ne. The average change due to genetic drift is no change -- the most likely result of random sampling is no change at all. Populations can change in either direction (larger or smaller) due to random sampling, and larger amounts of change are increasingly less likely. More change is likely in smaller populations, so that the amount of change depends on the effective population size. Lande gives an expression for the effective population size N* at which the observed amount of change is at the 95 percent confidence limit:

    If we assume h2 = 0.5, t = 25,000 generations, and the standard deviation is 1, then N* is estimated as 12,000. Since the effective population size of all extant hominoid species is around 10,000, this estimate is fully consistent with the evolution of P3 length by genetic drift alone.

    In fact, it's pretty hard to find anything in human evolution that couldn't have evolved by drift alone, under these assumptions. For example, Wolpoff and I (2001) found that the Middle Pleistocene increase in cranial capacity was consistent with genetic drift in a population with Ne = 1.8 x 106. The increase from early Neandertals to Würm Neandertals was less likely to occur by drift alone -- our estimate of Ne = 1.2 x 103 is quite a bit less than 10,000. On the other hand, there would be many who would argue that the effective population size in Europe alone really was that small, and that therefore the Neandertals increased their cranial capacity by genetic drift also.

    Now, the increase in endocranial volume in humans is one of the most impressive long-term evolutionary trends in mammals. If even that is explicable by genetic drift, then it is pretty clear that we don't ever need natural selection at all.

    So I should really like genetic drift, right? I mean, it explains everything, doesn't it?

    The fallacy

    Of course, what is actually going on is that we have chosen a null hypothesis that is especially hard to refute. This means we should expect a lot of type II error: using this method, we can't reject the hypothesis of genetic drift even if it isn't the right answer.

    What is worse, even if we were to show that the amount of phenotypic change is too great for a given effective population size, there will always be someone to argue that the effective population size was smaller in the past. So genetic drift is a moving target -- it is effectively impossible to reject.

    The operative problems here are (i) a relatively small amount of change over (ii) a very long period of time. This combination will usually be consistent with Lande's derivation for genetic drift and reasonable effective population sizes -- particularly if we cannot establish in advance what effective population size is actually reasonable.

    There is a big contrast between a long timescale and a short timescale in this comparison. Natural selection is certainly much faster than genetic drift on a short timescale -- the time to fixation of an adaptive allele by selection proceeds as the logarithm of population size, while the fixation time by drift proceeds linearly with population size. Genetic drift can change a population quickly, but only if the population is very small. Selection can change a large population quickly.

    The fallacy is the assumption that the difference between selection and drift over short timescales also is a difference over long timescales. There is some ultimate limit on the evolution of any character. Selection may make a mouse the size of a cat in a few hundred generations, but even assuming that those cat-sized mice can stick around, there is no reason to think that dog-sized mice will be better! At some point, selection will stabilize -- and for most characters the amount of change permitted by stabilizing selection is not too great. Sampled at time intervals of many hundreds of generations, selection may look exactly like genetic drift.

    What to do

    People think about genetic drift because of its mathematical convenience. Sampling error is predictable, and the repeated occurrence of sampling error over many generations follows well-known probability distributions.

    Selection is predictable too, but it requires you to actually know something about ecology. We often don't know anything, and when we do, we usually have some particular relationship in mind, which needs to be tested. So, we test the hypothesis of neutrality, with all its mathematical simplicity.

    But remembering that the null hypothesis is sometimes -- maybe even often -- true doesn't mean that we should be satisfied with any particular test of that null. For neutral evolution of phenotypes, there are more powerful tests than evolutionary rates. The problem is that these "tests" are not in large part quantitative, but instead are logical or qualitative.

    For example, it is very likely that human brains increased in size under selection because there should clearly have been selection against larger brains because of their energetic costs. The rate of evolution does not factor in here, and is in fact irrelevant to the assessment of selection.

    The case does require a more complex model than simple directional selection -- instead it involved a structured model in which the force of selection is mostly stabilized by a counterforce of selection. It also begs an explanation for why the change should have proceeded at a given rate -- for example, was it slow because of environmental constraints? Such constraints would seem a likely explanation for the rate of change of dental size in Australopithecus afarensis.

    But when most people talk about genetic drift, their reality doesn't seem to include the mathematical consequences of genetic drift.

    For one thing, genetic drift is sloooooow. It affects allele frequencies on a time scale in generations proportional to the effective population size.

    We sometimes hear that it is a bad thing to doubt the power and ubiquity of genetic drift. This doubt is sometimes equated with adaptationism, taken as the uncritical assumption of selection as a null hypothesis.

    References:

    Gould SJ, Lewontin RC. 1979. The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proc R Soc Lond B 205:581-598.

    Lockwood CA, Kimbel WH, Johanson DC. 2000. Temporal trends and metric variation in the mandibles and dentition of Australopithecus afarensis. J Hum Evol 39:23-55.

    Hawks J, Wolpoff MH. 2001. The accretion model of Neandertal evolution. Evolution 55:1474-1485.

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Neandertals

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

Denisova

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

Acceleration

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

Malapa

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