Koobi Fora

In case you haven't been paying attention, the chronology of early African Homo has been completely turned upside-down this year. Well, "upside-down" isn't precisely right; "displaced younger by a quarter-million years" is better.

The redating has come from Frank Brown's group, which in a series of papers has defined and dated stratigraphic units between the major tuffs of the Koobi Fora formation, between the KBS Tuff at 1.87 Ma and the Chari tuff at around 1.38 Ma. Gathogo and Brown (2006) outline the consequences of this redating for fossils of early Homo. Their paper focuses on the fossils from area 123 at Koobi Fora, but discusses the likely consequences of redating on other localities.

Fossils of Homo now estimated to be 1.65 +/- 0.15 myr in age in the Koobi Fora region are currently assigned to at least two taxa on the basis of both crania and mandibles. Homo habilis is represented by specimens KNM-ER 1501, 1502, 1805, and 1813, and H. ergaster is represented by specimens KNM-ER 730, 1812, and 3733 (for attributions, see Wood, 1991, 1992; Wood and Richmond, 2000). The ages of specimens KNM-ER 1501, 1502, 1812, and 1813 have been discussed above, and although not the main focus of this paper, a few notes are offered below on the others.

Specimen KNM-ER 730 derives from a level 5 m below the Koobi Fora Tuff Complex in Area 103 (Feibel et al., 1989), and is thus ca. 1.6 myr old. Feibel et al. (1989) gave an age of 1.85 myr for KNM-ER 1805, but this specimen lies "just below the base of the Okote Tuff" in Area 130 (Leakey et al., 1978), and is more likely closer in age to that of the base of the Okote Tuff Complex (ca. 1.6 myr) than it is to that of the KBS Tuff (1.87 myr). On the basis of mollusc-packed sandstones and algal horizons correlated from Area 102 to Area 104, Feibel et al. (1989) estimated that KNM-ER 3733 was 1.78 myr in age. Although the age of KNM-ER 3733 cannot be confirmed without additional fieldwork, the White Tuff, with an estimated age of 1.63 myr (Brown et al., 2006), is the nearest unequivocally identified unit in the local section in Area 104. This tuff is exposed Indeed, all specimens from Koobi Fora assigned to H. aff. H. erectus by Wood (1991), many of which are now referred to H. ergaster (Wood and Richmond, 2000), are now estimated to be 1.45 to 1.65 myr old with the exception of KNM-ER 2598. The latter specimen, which is a partial occipital bone from Area 15, was placed 4 m below the KBS Tuff by Feibel et al. (1989) and estimated to be about 1.9 myr old. This age estimate is reasonable because strata do not extend more than 7 m above or below the KBS Tuff at the recorded location of KNM-ER 2598 (Gathogo and Brown 2006:7-8, emphasis added).

This raises a question: Just how much evidence is left for large-bodied H. erectus-like hominids earlier than 1.65 Ma?

Wood (1991) didn't diagnose postcrania, and Gathogo and Brown (2006) don't comment on them. At least KNM-ER 1808 would seem to fall under this umbrella, since Wood (1991) did diagnose that. But more important in bracketing the evolution of large body size is KNM-ER 3228, a hip bone previously dated to 1.95 Ma. It's pretty big for a human, let alone an australopithecine. On the other hand, McHenry and Coffing (2000) suggested that KNM-ER 3228 might belong to H. rudolfensis. To my eyes, this would make it a pretty big specimen compared with femora like KNM-ER 1472 and KNM-ER 1481, but who knows?

Another uncomforable fit in an H. rudolfensis would be KNM-ER 2598. It sure looks like a large-brained, thick-boned specimen. It doesn't look much like KNM-ER 1470. But then, maybe 1470 is the unusual specimen...

Gathogo and Brown (2006) take on directly the issue of KNM-ER 1470 and KNM-ER 1813. The two were formerly considered contemporaries at around 1.89 Ma, but now KNM-ER 1813 is only 1.65 Ma.

KNM-ER 1813, lateral view

The real offshoot of this is that there are no longer any early small-skulled habilines. The question of whether KNM-ER 1470 and KNM-ER 1813 were too different to belong to a single species has drawn a lot of ink, but it was always a non sequitur, because the two weren't the only crania to consider. The more interesting observation had been that Olduvai Gorge preserved only small-skulled habilines, while Koobi Fora had both small and large ones. This was not only a geographic problem but also a temporal one, since the Olduvai habilines were all relatively late (less than around 1.8 Ma) and the Turkana habilines were mostly earlier.

Now the situation has changed. The small Turkana habiline, KNM-ER 1813, is now contemporary with the Olduvai sample. There are no longer any small-skulled early Turkana habilines. KNM-ER 1805 makes sense as a male of the later, small-skulled sample because it is relatively small-brained but robustly built (e.g., with a sagittal crest). That leaves KNM-ER 1470, KNM-ER 1590, KNM-ER 3732, and KNM-ER 3735 as plausible habilines before 1.85 Ma.

This seems like a nice sample as a possible ancestor for both later large-bodied Homo and later habilines. Heck, Wood (1991) even wrote this in his description of KNM-ER 3735:

Some features (e.g. vault thickness) ally it with a Homo erectus-like hominid, but in other areas (e.g. the frontal) it is more like crania such as KNM-ER 1813, a conclusion endorsed by Walker (1987) and by Leakey et al. (1989). Tobias (1989) includes KNM-ER 3735 within H. habilis (Wood 1991:134-135).

What more could you ask of a common ancestor? But then if some of this ancestral population would be expected to resemble later H. erectus-like specimens, then why not KNM-ER 2598?

And what, exactly, would make such a population -- with its mixture of H. erectus-like and habiline-like features -- different from Dmanisi? The answer, of course, is KNM-ER 1470. It's still the odd one in this lineup. But then, it does have the largest brain in this set, which might help to explain the rounded occiput.

Looking at what is left in the early part of the sequence is certainly interesting, but just as interesting is how all the H. erectus-like specimens are all bunched together between 1.65 and 1.45 Ma. This is the time interval that already held KNM-WT 15000, KNM-ER 3883, and KNM-ER 42700, and is just older than OH 9. Now we can add KNM-ER 3733, KNM-ER 730, KNM-ER 1808, and KNM-ER 1821. Isn't this an interesting sample? Don't you wish we knew about the other postcrania?

It seems to me that the hypothesis that H. erectus-like hominids first appeared in Africa around 1.65 Ma has interesting archaeological consequences. This isn't long before the appearance of the earliest Acheulean, and it plausibly makes the Developed Oldowan-Acheulean sequence a correlate of this evolution.

It is markedly not coincident with the earliest such evidence in Asia. But that raises the Dmanisi question again, doesn't it?

References:

Brown FH, Haileab B, McDougall I. 2006. Sequence of tuffs between the KBS Tuff and the Chari Tuff in the Turkana Basin, Kenya and Ethiopia. J Geol Soc 163:185-204.

Gathogo PN, Brown FH. 2006. Revised stratigraphy of Area 123, Koobi Fora, Kenya, and new age estiamtes of its fossil mammals, including hominins. J Hum Evol (in press) DOI link

McDougall I, Brown FH. 2006. Precise ⁴⁰Ar/³⁹Ar geochronology for the upper Koobi Fora Formation, northern Kenya. J Geol Soc 163:205-220.

Wood B. 1991. Koobi Fora Research Project, Volume 4, Hominid Cranial Remains. Clarendon Press, Oxford.

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

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

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

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

Here's the abstract:

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

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

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

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

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

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

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

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

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

But Dennell and Roebroeks give motivations for examining some alternatives.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

National Geographic News also has an article about the paper.

References:

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

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

One study reported at the meetings brought to mind a growing literature on the sophistication of Pliocene archaeological assemblages, grouped as "Oldowan" or "pre-Oldowan." Most Oldowan tools lack a standardized form, and the assemblages are dominated by flakes, core tools with one sharp edge or point, and unmodified manuports or hammerstones. The lack of standardized tools (such as handaxes) lead easily to the conclusion that Oldowan technical assemblages are simple, so that anyone could make one who could chip rock.

But a number of recent sources illustrate that this perception isn't accurate. The Oldowan encompasses a variation of manufacturing complexity and other elements, but cannot be said to be merely rocks that have been carried or bashed together. The most interesting studies have examined the issue of raw material exploitation and the actual processes used to reduce stone into useful tools.

David Braun gave a presentation that relates to this topic at the 2005 Paleoanthropology Society meetings. In an analysis of the lithic remains at the Kanjera South site (Kenya), the group found that the intensity of the reduction sequence varied with source material.

Kanjera South, dating to around 2.2 million years, preserves one of the largest early records of material culture. According to Plummer et al. (2001:810):

Hominins at Kanjera South utilized a wider variety of lithic raw materials than found at most Oldowan sites, some of which (chert, quartz, quartzite) must have been transported from outside the immediate vicinity of the deposits since they are not present in the local clast population.

In the study by Braun et al. (2005), fine-grained quartzite, which had a nearest source outcrop a long distance away from the site, was reduced more intensively than locally available rock. This reduction involved more flakes taken per core, and occasional retouch. They hypothesized that this difference was due to the hardness of the quartzite and its ability to hold an edge, which makes tools made on quartzite potentially more useful. The hominids apparently recognized these qualities in the raw materials and exploited the quartzite. We can probably infer that the transport distance was a result of this recognition as well.

Stout and colleagues (2005) report on the use of raw materials at Gona, which is at present the earliest site at which stone tools are found, dating to around 2.6 million years. This study sampled unmodified stone taken from conglomerates that represent natural accumulations. If stone were taken merely from local sources without selectivity, the modified stone assemblages from the site would be expected to match these conglomerates. As described by Stout et al. (2005:366-367), the past understanding of Oldowan toolmakers has suggested that they may not have exhibited a clear understanding of the technical qualities of their raw materials:

At Koobi Fora, the high frequency of basalt in local conglomerates (i.e. ancient stream channels) is closely mirrored by its predominance in the artifact assemblages (Toth, 1985; Schick, 1987; Isaac et al., 1997). Although quartz, chert and glassy volcanics are "reasonably easily available" in the conglomerates, these material types do not appear to have been specifically selected for (Isaac et al., 1997:268).

This paper gives an excellent review of the history of raw material choice. In brief, the literature review indicates that early toolmakers at Koobi Fora may have exercised some degree of selectivity by avoiding certain kinds of stone (vesicular lavas and weathered cobbles). By Olduvai Bed II (after 1.7 million years ago), some toolmakers were clearly going out of their way to acquire certain stone materials:

The increased use of chert at Olduvai around 1.65 Myr was clearly occasioned by the temporary exposure of rich sources of this material by the retreating waters of the paleo-lake (Hay 1976). Hominid toolmakers at this time readily appreciated the superior flaking properties of chert, leading to the formation of the earliest known special-purpose quarry site at MNK CFS (Stiles et al. 1974; Stiles 1998) (Stout et al. 2005:367).

So when did this technical knowledge really originate? Was it a skill that developed gradually over the million years between Gona and Olduvai Bed II?

Stout and colleagues find that the materials used for tools at Gona do not match the natural cobbles that were avaiable in a number of respects. In essence, the hominids ignored many kinds of stone that had fracture patterns or other properties that made them less suitable as raw materials. They (2005:368) state the conclusion most clearly at the end of their introduction:

The strong pattern of raw material selection seen at Gona demonstrates that low levels of selectivity are not universal in the Oldowan. The fact that the assemblages from Gona are the oldest known in the world also argues against an overarching temporal trend toward increasing selectivity within the Oldowan.

The flaking skill of early hominids is the subject of a paper by Roche and colleagues (1999) in Science. At the two localities of Lokalalei 1 and 2C, both dating to 2.34 million years ago on the west shore of Lake Turkana, this study accomplished refits of stone flakes and cores to directly inspect the reduction sequence. They found that this sequence was quite complicated in many of the instances of manufacture at the two localities:

At LA2C, the dominant reduction sequence consists of these more reduced cores. Unidirectional or multidirectional removals are flaked on a single debitage surface, from natural or prepared platforms. This knapping system allows the production of a long series of removals without changing the volumetric structure of the core. The repeated application by the knappers of the same technical principles to a whole series of cores, and during the reduction of each core, indicates an elaborate debitage scheme, implying motor precision and coordination. These principles include an appreciation of the quality of the collected raw materials, a judicious exploitation of the natural morphology of the blocks and the maintenance of adequate flaking angles during the entire debitage sequence. These show that the notion of production was already assimilated by a group of hominids in this particular area. This notion is integrated within a real debitage strategy, here well-mastered and unprecedented for this time period (Roche et al. 1999:59).

Upon this description and analysis of the debitage, the study bases the following conclusion:

Overall simplicity and similarities between assemblages are the two main arguments recently put forward to substantiate a technological stasis hypothesis between the 2.6 and 1.6 Myr time periods, and to merge the related assemblages into a single vast 'Oldowan' technocomplex. The stasis hypothesis cannot hold out against the detailed technological analysis of the LA2C lithic assemblage. There can be no doubt about the elaborate character of the LA2C debitage schemes, which are far more sophisticated than at any other Pliocene site (Roche et al. 1999:59).

In total, these studies attest to the technological expertise of some early toolmakers. It is perhaps important to remember that the archaeological record should be expected to reflect an average technical competence much less than that possible (or possibly even typical) of early hominids. For example, every instance of stone flaking by an expert toolmaker certainly was preceded by novice flaking by the same individual earlier in his life. Considering the early mortality evidenced by australopithecine (and early human) remains, many potential toolmakers must have died before reaching proficiency. And toolmaking expertise was probably not uniformly distributed among social groups, so that some individuals had the opportunity to learn by observation some relatively complex reduction sequences and material properties, while others would have had to acquire the same knowledge by long trial and error, if at all.

In this framework, later changes in the archaeological record may reflect social and life history changes as much as ecological changes or brain evolution. Toolmaking intensifies later in the Oldowan, raw material selectivity increases, and some standard forms and more technically sophisticated artifacts, such as bifaces, routinely occur. The increased mental complexity of Pleistocene hominids may be a cause, but the behavioral changes have no obvious anatomical, life history, or other correlate. The apparent sophistication of the earliest toolmakers argue against the hypothesis that the later Oldowan represents a fundamentally more intelligent hominid species, with specialized adaptations to tool manufacture lacking in earlier hominids. Instead, I would suggest that later Oldowan hominids had on average a greater retention of cultural knowledge as a function of more stable groups and a longer life history than australopithecines. This shift might or might not have been accompanied by an increase in the capacity to learn, as might be evidenced by a greater length of the juvenile growth period.

From a conceptual perspective, there is necessarily a real limit on the standardization that is possible within any "technocomplex," and particularly one that spans a million years or longer. The kind of information that is ultimately transmissible (either horizontally or vertically) among hominids over this extreme time period is of the most rarified form imaginable. A model of early archaeological assemblages that incorporated the probable manner and form of information transfer among groups (along with their global demography) would predict both a global lack of complexity and a great extent of variability within that simple context. The evidence of technical skills therefore may inform directly about the demography of groups and their relationships with each other. The extent to which such variability may be consistent with cultural diffentiation among groups in other primate species (such as chimpanzees) I take to be a testable hypothesis.

References:

Most references within the featured articles below:

Plummer T, Ferraro J, Ditchfield P, Bishop L, Potts R. 2001. Late Pliocene Oldowan excavations at Kanjera South, Kenya. Antiquity 75:809-810.

Roche H, Delagnes A, Burgal JP, Feibel C, Kibunjia M, Mourre V, Texler PJ. 1999. Early hominid stone tool production and technical skill 2.34 Myr ago in West Turkana, Kenya. Science 399:57-60.

Stout D, Quade J, Semaw S, Rogers MJ, Levin NE. 2005. Raw material selectivity of the earliest stone toolmakers at Gona, Afar, Ethiopia. J Hum Evol 48:365-380.