Dietary divergence of robust australopithecines

I’m writing a review of the “robust” australopithecines, and I’m reminded of how drastically our understanding of these hominins has changed in just the past decade. Functional interpretations of the skull initially led to the common wisdom that these animals ate lots of hard foods, and had the jaws and teeth to cash the checks written by their diets.

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Comparison of a “gracile” (left) and “robust” (right) Australopithecus face, from Robinson (1954).

While anatomy provides evidence of what an animal could have been eating, there is more direct evidence of what animals actually did eat. Microscopic wear on teeth reflects what kinds of things made their way into an animal’s mouth, presumably as food, and so provide a rough idea of what kinds of foods an animal ate in the days before it died. Microwear studies of A. robustus from South Africa had confirmed previous wisdom: larger pits and more wear complexity in A. robustus than in the earlier, “gracile” A. africanus suggested more hard objects in the robust diet (e.g., Scott et al., 2005). A big shock came a mere 8 years ago with microwear data for the East African “hyper robust” A. boisei: molars had many parallel scratches and practically no pitting, suggesting of a highly vegetative diet (Ungar et al. 2008).

robust microwear

Microwear in A. boisei (blue) and A. robustus (red). Although they overlap mostly for anisotropy (y-axis), they are completely distinct for complexity (x-axis). Data from Grine et al. (2012) and skull diagrams from Kimbel et al. (2004).

Stable carbon isotope analysis, which assesses what kinds of plant-stuffs were prominent in the diet when skeletal tissues (e.g. teeth) formed, further showed that the two classically “robust” hominins (and the older, less known A. aethiopicus) ate different foods. Whereas A. robustus had the carbon isotope signature of an ecological generalist, A. boisei had values very similar to gelada monkeys who eat a ton of grass/sedge. GRASS!

robust isotopes

Stable carbon isotope data for robust australopithecines. Data from Cerling et al. (2013) and skull diagrams from Kimbel et al. (2004). Note again the complete distinction between A. robustus (red) and A. boisei (blue).

ResearchBlogging.orgWhile microwear and isotopes don’t tell us exactly what extinct animals ate, they nevertheless are much more precise than functional anatomy and help narrow down what these animals ate and how they used their environments. This highlights the importance of using multiple lines of evidence (anatomical, microscopic, chemical) to understand life and ecology of our ancient relatives.


Cerling TE, Manthi FK, Mbua EN, Leakey LN, Leakey MG, Leakey RE, Brown FH, Grine FE, Hart JA, Kaleme P, Roche H, Uno KT, & Wood BA (2013). Stable isotope-based diet reconstructions of Turkana Basin hominins. Proceedings of the National Academy of Sciences, 110 (26), 10501-6 PMID: 23733966

Grine FE, Sponheimer M, Ungar PS, Lee-Thorp J, & Teaford MF (2012). Dental microwear and stable isotopes inform the paleoecology of extinct hominins. American Journal of Physical Anthropology, 148 (2), 285-317 PMID: 22610903

Kimbel WH, Rak Y, & Johanson DC (2004). The Skull of Australopithecus afarensis. Oxford University Press.

Robinson, J. (1954). Prehominid Dentition and Hominid Evolution Evolution, 8 (4) DOI: 10.2307/2405779

Ungar PS, Grine FE, & Teaford MF (2008). Dental microwear and diet of the Plio-Pleistocene hominin Paranthropus boisei. PloS One, 3 (4) PMID: 18446200

The strange days of yore

Today is not like the good ol’ days. In many ways things have changed for the better. For instance, in the good ol’ days, many paleontologists would find fossils but let nary a soul examine them; today, you can download high quality 3D models of many important fossils from both East and South Africa, completely for free!

Robert Broom’s (1938) account of the discovery of the first Paranthropus (or Australopithecus) robustus is also a reminder of the strangeness of the bygone days of yore:

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Wait for it …

In June of this year a most important discovery was made. A schoolboy, Gert Terblanche, found in an outcrop of bone breccia near the top of a hill, a couple of miles from the Sterkfontein caves, much of the skull and lower jaw of a new type of anthropoid. Not realizing the value of the find, he damaged the specimen considerably in hammering it out of the rock. The palate with one molar tooth he gave to Mr. Barlow at Sterkfontein, from whom I obtained it. Recognizing that some of the teeth had recently been broken off, and that there must be other parts of the skull where the palate was found, I had to hunt up the schoolboy. I went to his home two miles off and found that he was at the school another two miles away, and his mother told me that he had four beautiful teeth with him. I naturally went to the school, and found the boy with four of what are perhaps the most valuable teeth in the world in his trouser pocket. He told me that there were more bits of the skull on the hillside. After school he took me to the place and I gathered every scrap I could find; and when these were later examined and cleaned and joined up, I found I had not only the nearly perfect palate with most of the teeth, but also practically the whole of the left side of the lower half of the skull and the nearly complete right lower jaw.

What a wild time – Broom hunts down poor Gert, barges into the school, then makes the kid show him where he hacked the skull out of the rock. Poor, poor Gertie.

Maybe it was a different Gertie, but surely the reaction was the same.

Maybe it was a different Gertie, but surely the reaction was the same.

Of course, there was a lot at stake. I mean, brazen Gert harbored not just “beautiful teeth,” but “the most valuable teeth in the world.” IN HIS TROUSERS! And of course Gert was also the soul possessor of priceless intel – the source of the fossils. So maybe Broom was justified in this zealous abduction. And O! such prose in a Nature paper! WAS IT WORTH IT, DR. BROOM?

At Sterkfontein, a bronzed Broom considers the weight of his actions.

At Sterkfontein, a bronzed Broom considers the weight of his actions.

Of course, Gert wasn’t the last kid to discover an important human fossil. The game-changing Australopithecus sediba  was discovered when Matthew Berger, son of famed Lee Berger and only 9 years old at the time, saw a piece of a clavicle sticking out of a block of breccia. Both Gert and Matthew show that you don’t have to be a doctor to make amazing discoveries. What future fossil discoveries will be made by kids, and making my adult accomplishments pale in comparison?!

Gracile & robust Australopithecus

Last week, I introduced my Human Evolution students to the “robust” australopithecines. It was a very delicate time, when we had to have a grown up, mature conversation about adult things. I reminded the students that they’re only human, but they must resist urges that seem only natural. No matter how much they want to, even if their friends are doing it, they must not act on the deep, dark desire to say that “robust” vs. “gracile” Australopithecus differ in their body build.

Don't do it, Homo naledi. Don't talk about body size when you mean to talk about jaw and tooth size. Illustration by Flos Vingerhoets.

Don’t do it, Homo naledi. Don’t talk about body size when you mean to talk about jaw and tooth size. Illustration by Flos Vingerhoets.

Every semester, students (who don’t read and/or pay attention to lecture) think that the difference between these two groups has to do with the species’ body sizes. This is a misconception that has reached the highest echelons of reference:

At least one person is not citing their source here. F-.

Apple and Google, at least one person here is not citing their source: F-. Also, is no one else surprised that this term is allegedly specific to anthropology?

No. In the case of australopiths, “gracile” and “robust” refer to the relative size of the jaws, teeth and chewing muscles (all contributing to the “masticatory apparatus”). Traditionally,  graciles include the ≥2 million year old Australopithecus afarensis and africanus, and robusts include the later A. boisei and robustus. The discovery of an A. aethiopicus cranium (Walker et al. 1986) somewhat blurred the lines between the two groups but it is usually included with the robusts (who are often collectively called Paranthropus). John Fleagle’s classic textbook (1999) illustrates the gracile-robust dichotomy very nicely:

Comparison of gracile (left) and robust (right) craniodental traits. From Fleagle, 1999.

So to recap: Jaws and teeth, people! To the best of my knowledge, there’s little or no evidence that the various australopithecines differed appreciably in body size (McHenry and Coffing, 2000), stoutness, or muscularity. Although the OH 80 partial skeleton, attributed to Australopithecus boisei  based on tooth size and proportions, includes a humerus with very thick cortical bone and a radius with a crazy big insertion for the biceps muscle – it was a very large and muscular A. boisei (Domínguez-Rodrigo et al., 2013). Nevertheless, gracile and robust australopithecine species differ most notably in their jaws and teeth, not bodies. Maybe this is why Liz Lemon was so confused about the term “robust”?

Today, these are somewhat antiquated terms. Back when the only hominins known to science were the species listed above, it was easy to make a distinction. However, as the fossil record has expanded of late, the gracile-robust dichotomy becomes blurry. Australopithecus garhi (Asfaw et al., 1999) has overall tooth proportions comparable to graciles, but absolute tooth sizes and sagittal cresting like robusts. The recently described Australopithecus deyiremeda has tooth sizes and proportions like graciles but lower jaws that are very thick, like those of robust australopithecines (Haile-Selassie et al., 2015).

So in light of all the confusion and blurring distinctions, maybe it’s time to scrap “gracile” vs. “robust”?

Further reading:  The “robust” australopiths (Constantino, 2013).

Asfaw B, White T, Lovejoy O, Latimer B, Simpson S, & Suwa G (1999). Australopithecus garhi: a new species of early hominid from Ethiopia. Science (New York, N.Y.), 284 (5414), 629-35 PMID: 10213683

Domínguez-Rodrigo, M., Pickering, T., Baquedano, E., Mabulla, A., Mark, D., Musiba, C., Bunn, H., Uribelarrea, D., Smith, V., Diez-Martin, F., Pérez-González, A., Sánchez, P., Santonja, M., Barboni, D., Gidna, A., Ashley, G., Yravedra, J., Heaton, J., & Arriaza, M. (2013). First Partial Skeleton of a 1.34-Million-Year-Old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania PLoS ONE, 8 (12) DOI: 10.1371/journal.pone.0080347

Haile-Selassie Y, Gibert L, Melillo SM, Ryan TM, Alene M, Deino A, Levin NE, Scott G, & Saylor BZ (2015). New species from Ethiopia further expands Middle Pliocene hominin diversity. Nature, 521 (7553), 483-8 PMID: 26017448

Walker, A., Leakey, R., Harris, J., & Brown, F. (1986). 2.5-Myr Australopithecus boisei from west of Lake Turkana, Kenya Nature, 322 (6079), 517-522 DOI: 10.1038/322517a0

Calotte or Carapace?

Is this the top of a hominid skull, replete with sagittal crest running down the middle, or is it the top of a tortoise shell?

This image comes from great resource I just found (thanks to Louise Leakey on Twitter) for paleoanthropology students – I won’t answer here whether this is hominid or turtle, you’ll have to find it at the African Fossils site.

The site has 3D, manipulable images of fossil hominids and other animals from Kenya and Tanzania. The Smithsonian Museum of Natural History also has a very nice 3D collection, similarly manipulable. Resolution isn’t always what you might want it to be (for instance, you won’t be able to tell if the basi-occipital suture is fused in the Homo erectus cranium KNM-ER 42700), but you still get good overall view of some neat and bizarre animals. Like this robust australopithecus! (KNM-ER 406) Hey, its brain case does look kinda like the pic above…

New beef with boisei – maybe the dingo ate their babies?

ResearchBlogging.orgUnfortunately, the title is not in reference to a study demonstrating that early hominids fell prey to wild dogs. But Elaine Benes would have appreciated it.

As part of my Latitudes Tour, I’m in Nairobi for a couple of days, hoping to spend some quality time with the young Australopithecus boisei kids at the Nairobi National Museum. Recall (that is, if I’ve mentioned it here?) that my dissertation research is on growth of the lower jaw, in Australopithecus robustus as compared to modern humans. The study of growth obviously requires analyzing individuals across different age groups (an “ontogenetic series” is the fancy term). Admittedly, then, the focus on A. robustus is chiefly because this species has the largest ontogenetic sample of any early hominid (tho at nearly 15 subadults, it’s still not as large as one could hope). Also because A. robustus was totally badass.

Australopithecus boisei makes an important comparison for A. robustus, because the two species are allegedly evolutionary ‘sisters’ – the “robust” australopithecines (though I’m personally not convinced that these two are each other’s closest relative). So their growth should be pretty similar. At the same time, though, A. boisei shows much greater adaptations to heavy chewing – they’ve been referred to as “hyper-robust.” So comparing growth in these species should elucidate how their differences arise.
Problem is, there just aren’t enough kids! It’s like that song by Arcade Fire. Wood and Constantino (2007) published a pretty comprehensive review of A. boisei, including a 1.5-page table of the skulls and teeth attributed to the species. So far as I know, only 4 specimens in this table are subadult mandibles, and so far as I can tell, they’re all about the same age (right around the age that the first permanent molar comes in). There are so many jaws of adult A. boisei (although many of these are abraded mandibular bodies lacking teeth) – so how can there be fewer subadults?!?!

A very preliminary observation of infant-child pairs in the two species suggests they both increase in size fairly dramatically between when they only have baby (a.k.a. “deciduous” or “milk”) teeth and when the first permanent molar comes in. But this is just a preliminary observation based on 2 specimens of each species! Take with a grain of salt!
On second thought, maybe I’ll propose the nearly untestable hypothesis that bone-eating hyenas ate the boisei babes, and that’s why we don’t have their jaws. What could have been nicely preserved subadult boisei bones are instead coprolites (fossilized poops). A little spectacular, yes, but it’s also been hypothesized that many of the A. robustus fossils we know and love came to us as carnivores’ scraps.
further reading:
Wood, B., & Constantino, P. (2007). Paranthropus boisei: Fifty years of evidence and analysis American Journal of Physical Anthropology, 134 (S45), 106-132 DOI: 10.1002/ajpa.20732

Culinary trends in an extinct hominid

A few weeks ago I discussed a recent paper that analyzed the carbon and oxygen isotope ratios from Australopithecus boisei molars (Cerling et al. 2011). The major finding here was that an enlarged sample (n=24 more) corroborated earlier isotopic (van der Merwe et al. 2008) and tooth wear evidence (Ungar et al. 2008) that A. boisei probably did not subsist on as much hard foods as previously thought. Although this strange hominid probably ate mostly grass/aquatic tubers, some researchers think it may have looked something like this:
Left, A. boisei reconstructed skull, from McCollum (1999, Fig. 1). Right, artist’s reconstruction of what the individual on the left may have looked like during life.
But looking at the numbers I’m wondering if the carbon isotopes reveal anything more about this curious hominid. If we plot boisei‘s carbon 13 values against the fossils’ estimated ages, there’s a small hint of a temporal trend, of increasing carbon 13 levels over time (more C4 plant consumption). Fitting a line to these data does indicate an increasing C4 component over time, but the slope of the line is not significantly different from zero. The early, high value could be an outlier (not eating the same stuff as his/her peers?), although the lowest carbon 13 value of all that would support this trend is also much lower than the other values; it could be a more anomalous one. So while it’s tempting to hypothesize dietary change over time in A. boisei, at the moment it looks like you can’t reject the hypothesis that diet is consistent throughout the Pleistocene until the A. boisei’s demise.  Supporting dietary stasis, Ungar and colleagues (2008) reported similar molar tooth wear in specimens from 2.27-1.4 million years ago.
In addition, Cerling and colleagues sampled at least one of each of the cheek teeth. Because teeth form in the jaws in a sequence (not all at the exact same time), the isotopic signatures from given teeth represent the dietary intake of carbon at various different points in an individual’s childhood. In the figure below I lumped upper and lower teeth together; the un-numbered “M” indicates molars unassigned to a specific position.

The first molar crown starts to form right around birth, and note here that it’s carbon 13 values are slightly higher than the other molars. The premolars and second molar start to form around the same time, so it is curious that each of these teeth show distinctly different ranges of carbon 13 levels. The sole P3 is also the lowest value (eating fewer C4 plants) in the entire sample, but the P4 has less negative values (eating more C4 plants). Not sure what’s going on here, but maybe later analyses of more specimens will clarify the situation.
Our australopithecine ancestors and cousins have proven to be a rag-tag bunch of funny bipeds, and A. boisei has proven to be one of the weirder ones, in my opinion. Of course descriptions of Ardipithecus ramidus and Australopithecus sediba skeletons have been recent reminders that we have lots left to learn about Pleistocene hominids. For my part, I’m interested in working out the deal with the group of “robust” Australopithecus.
Cerling, T., Mbua, E., Kirera, F., Manthi, F., Grine, F., Leakey, M., Sponheimer, M., & Uno, K. (2011). Diet of Paranthropus boisei in the early Pleistocene of East Africa Proceedings of the National Academy of Sciences DOI: 10.1073/pnas.1104627108
McCollum, M. (1999). The Robust Australopithecine Face: A Morphogenetic Perspective Science, 284 (5412), 301-305 DOI: 10.1126/science.284.5412.301
Ungar PS, Grine FE, & Teaford MF (2008). Dental microwear and diet of the Plio-Pleistocene hominin Paranthropus boisei. PloS one, 3 (4) PMID: 18446200
van der Merwe NJ, Masao FT and Bamford MK. 2008. Isotopic evidence for contrasting diets of early hominins Homo habilis and Australopithecus boisei of Tanzania. South African Journal of Science 104: 153-155

What the hell was Australopithecus boisei doing?

A little over 2 million years ago there a major divergence of hominids, leading on the one hand to our earliest ancestors in the genus Homo, and on the other hand to a group of ‘robust’ australopithecines, the latter group a failed evolutionary experiment in being human. In our ancestors, parts of the skull associated with chewing began to get smaller and more delicate, while the robust australopithecines increased the sizes of their crushin’-teeth and chewin’-muscle attachments.
A face not even a mother could love, so now they’re extinct (from McCollum 1999 Fig. 1). Note the very tall face, flaring cheeks, and massive lower jaw which would have facilitated wicked-pisser chewing power.
Weirder, there is a South African form (Australopithecus robustus) and an East African form (A. boisei, the figure here looks like it’s based off this species) of robust australopithecine. These two may have inherited their robust adaptations from a common ancestor, or they may be unrelated lineages that evolved these features in parallel. A boisei has been referred to as ‘hyper-robust,’ its face and teeth are generally larger than those of A. robustus.
For a while it’s been supposed that these ‘robust’ chewing adaptations in our weird, extinct evolutionary cousins (every family has those, right?) reflected a diet of hard objects requiring powerful crushing and grinding – things like hard fruits, seeds, Italian bread, etc. But a few years ago Peter Ungar and others (2008) examined the microscopic wear patterns on the surfaces of molar teeth of A. boisei and noted that they lacked the characteristic pits of a hard-object feeder. A. robustus on the other hand does have wear patterns more like an animal that ate hard foods. Why such a difference? Why the hell wasn’t boisei behaving robustly?
Also in 2008 Nikolaas van der Merwe and colleagues analyzed the carbon isotopes preserved in the teeth of A. boisei and some other fossils. Briefly, plants utilize two isotopes of carbon (C12 and C13), but ‘prefer’ the lighter-weight C12. Some groups of plants like grasses have thrived because they’re less picky and can get by just as well with C13. Different kinds of plants, then, incorporate different amounts of these two carbon isotopes into their tissues, then when animals eat it, these isotopes get incorporated into the animal’s developing tissues, including tooth enamel. So by looking at the relative amounts of carbon in teeth, researchers can get a rough idea of whether an animal was eating more of the C13-loving or C13-loathing plants (or the animals eating the plants). van der Merwe and others found A. boisei to have a way higher percentage of the plants that don’t discriminate against C13 as much, possibly things like grass, sedges or terrestrial flowering plants. GRASS?!

Last week, Thure Cerling and colleagues expanded on the earlier study led by van der Merwe, including a larger set of boisei specimens spanning 500 thousand years of the species’ existence. Lo and behold, Cerling and others got similar results: the isotopic signature in A boisei is similar to grass-feeding pigs and horses in its habitat – was the badass “hyper robust” A boisei just a hominid version of a horse? Now, the silica in grass make it extremely wearing on tooth enamel, and while A. boisei had crazy thick molar enamel, I would be a little surprised if the boisei dentition could withstand a lifetime of a grassy diet. Nevertheless, boisei‘s diet clearly differed from robustus, based on both dental wear and carbon isotopes.
This raises interesting questions about the evolution of the robust group. Does their shared ‘robust’ morphology reflect common ancestry, with the subtle differences the result of their divergent diets? Or do the subtle differences indicate that they evolved separately but their diets for whatever reasons resulted in similar mechanical loading on their jaws and faces? It should also be noted that while the dates for South African cave sites are always a bit uncertain, it is possible that A. robustus persisted alongside genus Homo until around 1 million years ago, whereas the fossil record for A. boisei craps out around 1.4 million years ago – was A. boisei too specialized on crappy grass, resulting in its evolutionary demise?
A horse-ish, human-ish hominid? Australopithecus boisei, rest in peace. 2.1 – 1.4 mya.
Cerling TE, Mbua E, Kirera FM, Manthi FK, Grine FE, Leakey MG, Sponheimer M, & Uno KT (2011). Diet of Paranthropus boisei in the early Pleistocene of East Africa. Proceedings of the National Academy of Sciences of the United States of America PMID: 21536914
McCollum, M. (1999). The Robust Australopithecine Face: A Morphogenetic Perspective Science, 284 (5412), 301-305 DOI: 10.1126/science.284.5412.301
Ungar PS, Grine FE, & Teaford MF (2008). Dental microwear and diet of the Plio-Pleistocene hominin Paranthropus boisei. PloS one, 3 (4) PMID: 18446200
van der Merwe NJ, Masao FT, & Bamford MK (2008). Isotopic evidence for contrasting diets of early hominins Homo habilis and Australopithecus boisei of Tanzania. South African Journal of Science 104: 153-155

As promised, malaria resistance in baboons

Last week I started to mention a recent paper in Nature on the evolution of malaria resistance in baboons, but then went out and partied instead. Not wanting to be a bastard, I’d better make good. While I’ll try to pull a good lesson from this, be warned that I’m about to discuss a topic about which I am no expert.

Malaria sucks, you don’t want to get it. There are anti-malaria medications out there, but I understand that they can make you insane, or at least have crazy dreams. Fortunately for millions of humans, there is a genetic basis for malaria resistance, so they don’t have to buy the anti-malaria crazy pills. Now, the paper tells me that a polymorphism in part of the FY gene turns the gene off in red blood cells, and that individuals with this variant are then strongly protected from malaria. No Lariam for these folks.

Jenny Tung and colleagues analyzed the homologous region of the FY gene in almost 200 yellow baboons (Papio cynocephalus) from Kenya, as well as tested these baboons for Hepatocystis parasites–relatives of Plasmodium vivax, which don’t cause malaria in baboons, but does really suck for them. And wouldn’t you know it–this same region on the baboon FY gene is also associated with Hepatocystis infection, where individuals with certain genetic variants have a lower susceptability to infection!

Now, the underlying genetic architecture and subsequent mechanisms of infection resistance are not exactly the same. But here’s the take home message from the paper:

“These results suggest that the genetic basis of phenotypic variation in different primate species can exhibit a remarkable degree of parallelism. In this case, not only are the similarities present at the molecular level . . . but they also extend to the mechanism that links molecular and phenotypic variation”

In other words, closely related species are equiped with very similar (or often the same) genetic or developmental “hardware,” and so evolution can cause them to come up with similar solutions to the same problem. In this case, there’s a similar genetic basis underlying infection resistance in humans and baboons. But I think this is a lesson that can be extended to, or at least kept in mind when considering, phenotypic evolution generally.

I’ve always (well, for the past three and a half years since I’ve been studying physical anthropology) thought that such a situation might characterize the “robust” australopithecines of East and South Africa. It is possible that these groups are not each others’ closest relatives, but that they evolved many craniodental characters in parallel, in response to selection for a heavy-chewing diet. This becomes even more plausible if it should turn out that many of these cranial and dental features are morphologically integrated–something I’m working on at the moment (if anyone reads this and scoops me, you will pay).

So, interesting paper. Reference

Tung, J. et al. Evolution of a malaria resistance gene in wild primates. Nature, in press.

Zach, get over the robust australopithecines

Let’s return to my favorite group (though I think they’re not terribly closely related) in the human fossil record, the robust australopithecines. They have popped fairly frequently in the news this year, most recently regarding their possible use of bone tools <!–[if supportFields]> ADDIN EN.CITE Backwell21221217Backwell, Lucindad’Errico, FrancescoEarly hominid bone tools from Drimolen, South AfricaJournal of Archaeological ScienceJournal of Archaeological ScienceIn Press, Accepted Manuscript <![endif]–>(Backwell and d’Errico, in press)<!–[if supportFields]><![endif]–>. This group is funny. They first appeared probably some time around 2.7 million years ago, in the controversial form of Australopithecus aethiopicus. No one knows where exactly this enigmatic group came from, save for that the only known, fairly complete cranium (KNM WT 17000) has many primitive features, and is largely similar to the earlier A. afarensis. Around 2.3 million years ago or so, A. aethiopicus appears to be ‘replaced’ by A. boisei, whose face is less protruding, has smaller anterior teeth, and has a derived P3 morphology. This is all in East Africa, mind you. Then, some time probably around 2 million years ago, or a little later, a robust form (A. robustus) appears in South Africa, where erstwhile the only hominin was A. africanus (some argue that there are 2 species in the A. africanus hypodigm). Personally, A. robustus looks like a more ‘robust’ A. africanus: larger posterior teeth, more anteriorly placed cheeks—but there is much overlap in many of traits between these two taxa. And much to the chagrin of many cladists, the South and East African robusts appear fairly different morphologically; a recent study <!–[if supportFields]> ADDIN EN.CITE Gonzalez-Jose200821021017Gonzalez-Jose, RolandoEscapa, IgnacioNeves, Walter A.Cuneo, RubenPucciarelli, Hector M.Cladistic analysis of continuous modularized traits provides phylogenetic signals in Homo evolutionNatureNature775-77845371962008Nature Publishing Group <![endif]–>(Gonzalez-Jose et al. 2008)<!–[if supportFields]><![endif]–> that examined hominoids morphometrically (that is, in terms of aspects of cranial shape) found the two robust taxa to be distinct (but that’s a topic for another post . . .). And all the while these buggers lived right alongside Homo, our ancestors! That’s some effed up stuff.

Now for the recent paper. Blackwell and d’Errico (in press) report on an assemblage of bone tools from the site of Drimolen in S. Africa (~2-1.5 million years ago). Drimolen is very near Swartkrans and Kromdraai, two other cave sites with a wealth of A. robustus, and to a lesser extent Homo, material has been recovered. First, how do they know these bones were tools? The tools were compared to other bones, known to have been worn (down) by other processes, like gnawing or carnivore chewing, and the tools appear quite different from these. Also, experimental studies of actually using bones as digging tools (to dig up underground tubers and especially to dig into termite mounds) have produced the same kind of wear as the fossil bone tools. Finally, many bone tools are known from the site of Swartkrans, which as I mentioned above is also laden with A. robustus remains. So it appears pretty likely that A. robustus (or Homo) was using bone tools to dig for foodstuffs around Drimolen and other cave sites in the early Pleistocene. *Note: our friend and colleague Julie Lesnik is currently finishing up her fieldwork, examining bone tools from Swartkrans, and observing chimpanzees digging for termites in Senegal; perhaps we could get her to write a good post on the topic.

Second, how do they know who used these tools? This an excellent question, which similarly plagues the postcranial material from these S. African sites (and even E. African sites, cf. the OH 7 hand and the OH 8 foot, again another topic for another post on another day . . .). The basic argument, similar to that for the postcrania, is that where these bone tools are found, the hominin assemblage is dominated by A. robustus remains; well, at least there are more A. robustus teeth and cranial material relative to comparable Homo fossils. So it’s guilt by association for the bone tools and postcrania here. Of course the only real way to test the hypothesis that most S. African postcrania and bone tools are to be associated with A. robustus and not Homo would be to find a complete skeleton of either, and to find bone tools in strong association (i.e. almost still in the clutches) of a certain taxon. Another argument used in support of robustus as the bone tool user is the fact that stone tools—the hallmark of early humankind?—are relatively absent at these sites. So it’s probable, but certainly not unequivocally proven, that A. robustus was the user of these stone tools (note that the authors never make any claim of such proof).

The authors use the bone tool assemblages from Swartkrans and Drimolen to infer ‘cultural’ behavior patterns for A. robustus. A recent paper claimed that since larger (male) A. robustus cranial specimens tended to be older than smaller ones, that this was evidence for extended male growth in the taxon <!–[if supportFields]> ADDIN EN.CITE Lockwood20079917Lockwood, Charles A.Menter, Colin G.Moggi-Cecchi, JacopoKeyser, Andre W.Extended male growth in a fossil hominin speciesScienceScience1443-1446318Australopithecus robustusdelayed maturation200730 November 2007doi:10.1126/science.1149211<![endif]–>(Lockwood et al. 2007)<!–[if supportFields]><![endif]–>. This, coupled with the pronounced sexual dimorphism in this taxon (and all hominins until only fairly recently), suggested Lockwood et al. that A. robustus had a social structure similar to that of gorillas, in which one or two males associate with a number of females and their offspring (a simpler answer is that selection favored larger males, and so they were living longer, but Mary and I had trouble trying to demonstrate this). So Blackwell and d’Errico figure that this evidence, along with data from chimpanzee termite-foraging behavior, suggests that females were the predominant practitioners of termiting using bone tools.

Why do I think this is interesting? I think this shows a potentially very important ecological divergence between the robust australopithecine lineages on the on hand, and between A. robustus and Homo on the other. Could the disparate toolkits of these hominins have played an important part in the evolution of these lineages? It is debated whether robust australopithecine hands were physically capable of actually making stone tools (which is quite difficult). Part of this stems from the fact that it’s hard to find a complete hand (damned beetles…) and then to attribute it to a specific taxon. For instance, the OH7 partial hand, which was supposed to be part of the H. habilis holotype, was recently found to be most similar morphologically to A. robustus from S. Africa, and functionally not adapted to tool-making <!–[if supportFields]> ADDIN EN.CITE Moyà-Solà200826126117Moyà-Solà, S.Köhler, M.Alba, D. M.Almécija, S.Taxonomic Attribution of the Olduvai Hominid 7 Manual Remains and the Functional Interpretation of Hand Morphology in Robust AustralopithecinesFolia PrimatologicaFolia Primatologica215-2507942008 <![endif]–>(Moyà-Solà et al. 2008)<!–[if supportFields]><![endif]–>. Of course, this relies on assumptions about whether hand material from Swartkrans represents A. robustus or Homo. Nevertheless, it may well be that part of the adaptive divergence of A. robustus and Homo was the former’s use of bone tools to dig for termites, while the latter was able to manipulate stone to exploit higher quality resources. Cool.

Also significantly, I think this points to more evidence against uniting A. robustus and boisei into the genus Paranthropus. Yes, they both had large posterior teeth (though boisei’s were generally larger) and faces built to house large chewing muscles. But other than that they appear pretty different. As noted above, a recent study looking at modularized traits in crania found that the two robust groups were not monophyletic, which is a criterion when making taxonomic decisions (Gonzalez-Jose et al. 2008). A. robustus has striking affinity with the earlier A. africanus, while A. boisei has striking affinity, to the exclusion of A. robustus, with A. aethiopicus, suggesting that A. robustus and boisei didn’t share a common ancestor. It has also been argued that sharing an ecological space is another criterion for generic membership <!–[if supportFields]> ADDIN EN.CITE Wood199926226217Bernard WoodMark CollardDepartment of Anthropology, George Washington University, 2110 G Street NW, Washington, DC 20052, USA; Human Origins Program, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA.; Department of Anthropology, University College London, Gower Street, London WC1E 6BT, UK.The changing face of genus HomoEvolutionary AnthropologyEvol AnthropolEvolutionary Anthropology195-2078619991520-6505;2-2 10.1002/(SICI)1520-6505(1999)8:63.0.CO;2-2<![endif]–>(Wood and Collard 1999)<!–[if supportFields]><![endif]–>. But isotopic and dental microwear evidence show A. robustus and boisei to have been distinct. Also, (shame on me) I’m not sure the extent to which A. boisei is associated with bone tools, but to my knowledge it is not. So here we have morphological, geological, temporal, and ecological discontinuity between the two main ‘robust’ taxa. That pretty much sinks Paranthropus for me. I don’t know why I am so adamant about these taxa’s paraphyly, but I am. I feel that taxonomy should reflect important biological and phylogenetic reality. And it seems to me that because there isn’t compelling evidence that A. robustus and boisei share the same ancestor, or interbred, that they shouldn’t be taxonomically separated from the other australopithecines. And that’s my two cents.


Backwell L, d’Errico F Early hominid bone tools from Drimolen, South Africa. Journal of Archaeological Science In Press, Accepted Manuscript

Gonzalez-Jose R, Escapa I, Neves WA, Cuneo R, Pucciarelli HM (2008) Cladistic analysis of continuous modularized traits provides phylogenetic signals in Homo evolution. Nature 453(7196):775-778

Lockwood CA, Menter CG, Moggi-Cecchi J, Keyser AW (2007) Extended male growth in a fossil hominin species. Science 318:1443-1446

Moyà-Solà S, Köhler M, Alba DM, Almécija S (2008) Taxonomic Attribution of the Olduvai Hominid 7 Manual Remains and the Functional Interpretation of Hand Morphology in Robust Australopithecines. Folia Primatologica 79(4):215-250

Wood B, Collard M (1999) The changing face of genus Homo. Evolutionary Anthropology 8(6):195-207<!–[if supportFields]><![endif]–>

New twist from teeth

Peter Ungar, Fred Grine and Mark Teaford recently reported in PLoS ONE on their results of studying the microwear on Australopithecus boisei molars. Their study showed that the microwear differs from that of A. robustus, arguably boisei‘s South African counterpart, and from A. africanus. Here’s the abstract:

The Plio-Pleistocene hominin Paranthropus boisei had enormous, flat, thickly enameled cheek teeth, a robust cranium and mandible, and inferred massive, powerful chewing muscles. This specialized morphology, which earned P. boisei the nickname “Nutcracker Man”, suggests that this hominin could have consumed very mechanically challenging foods. It has been recently argued, however, that specialized hominin morphology may indicate adaptations for the consumption of occasional fallback foods rather than preferred resources. Dental microwear offers a potential means by which to test this hypothesis in that it reflects actual use rather than genetic adaptation. High microwear surface texture complexity and anisotropy in extant primates can be associated with the consumption of exceptionally hard and tough foods respectively. Here we present the first quantitative analysis of dental microwear for P. boisei. Seven specimens examined preserved unobscured antemortem molar microwear. These all show relatively low complexity and anisotropy values. This suggests that none of the individuals consumed especially hard or tough foods in the days before they died. The apparent discrepancy between microwear and functional anatomy is consistent with the idea that P. boisei presents a hominin example of Liem’s Paradox, wherein a highly derived morphology need not reflect a specialized diet.

Note that they refer to boisei and robustus as “Paranthropus,” whereas I (and others) refer to them as Australopithecus. A. boisei and robustus are two “robust” australopithecines, described as such because their skulls and teeth suggest these guys were adapted for prolonged, powerful bouts of mastication (it means chewing, get your mind out of the gutter). Some people argue that these two taxa form a monophyletic group; that is, they share a last common ancestor that is not shared by any other taxon. If this is the case, the generic distinction (Paranthropus) can be made, separating them from the other australopithecines. Though I tend to lump groups, I really think that these taxa do not form a monophyletic group, that they have different ancestors (that their superficially similar masticatory apparati were independently evolved), and that they should stay in the genus Australopithecus. Right now, this issue (wherein I am very interested) has yet to be resolved.

Anywho, what’s important here is that the two robust austrlopithecines differ in their microwear patterns, which suggests that the two subsisted on different diets. Similarly, Wood and Constantino (2007) report that the stable carbon isotope signal from boisei (yet unpublished, but communicated to them personally by Matt Sponheimer) is different from the A. robustus and africanus. Together, these two data indicate that the robust australopitheciens (not to speak about A. aethiopicus…) were quite different in their diets (and possibly lifestyles?). Interestingly, A. robustus‘s molar microwear and stable isotope signals are very similar to that of A. africanus, who was present in the same regions as robustus but a bit earlier in time. This bolsters the scenario in which A. robustus is evolved from A. africanus, or something like it. Could this suggest also that A. boisei is not descendant from A. africanus? Or, is it simply that there were different foods available in the Plio-Pleistocene of South and East Africa?

Another important note that the authors bring up is the fact that of the seven specimens examined, none appeared to have eaten tough or hard foods that might necessitate the use of their (we assume) powerful masticatory muscles. Now why the hell would they have such a derived face, jaws and teeth if they were not eating things that would have required such an apparatus? One proposed scenario about the “hyper-robust” masticatory apparatus of A. boisei is that it is an adaptation for only the toughest of times, when survival might have hinged upon the ability to process and ingest the lowest quality (and hardest to eat) foods. Ungar et al.’s data suggest that this may well be the case, that the powerful masticatory apparatus came in handy only very rarely, and so the dietary signal from microwear reflects what these critters usually ate (and preferred to eat).

If this is really the case, then it might suggest that the robust face of boisei was almost completely genetically acquired, that epigenetic factors did not contribute greatly to produce boisei‘s face. This could be important for teasing out criteria (i.e. skeletal, craniofacial traits) useful in phylogenetic reconstruction. For example, it could be that certain robust features of boisei‘s face indicate a shared genetic ancestry, whereas those of robustus were more epigenetic in nature, acquired over a lifetime of experiencing high chewing forces. Contrariwise, these traits might be the result of these two taxa’s shared ancestry.

Either way, this paper presents interesting new information about the most bizarre hominin evolutionary dead-end, the robust australopithecines.


Ungar PS, Grine FE, and Teaford MF. 2008. Dental Microwear and Diet of the Plio-Pleistocene Hominin Paranthropus boisei. PLoS ONE 3(4):e2044.

Wood B, and Constantino P. 2007. Paranthropus boisei: Fifty years of evidence and analysis. American Journal of Physical Anthropology 134(S45):106-132.