#FossilFriday: 2015 Retrospecticus

January 1, 2016January 1, 2016 / zcofran / Leave a comment

Holy crap 2015 was a big year for fossils. And how fortuitous that 2016 begins on a Fossil Friday – let’s recap some of last year’s major discoveries.

Homo naledi

Some Homo naledi mandibles in order from least to most worn teeth.

The Homo naledi sample is a paleoanthropologist’s dream – a new member of the genus Homo with a unique combination of traits, countless remains belonging to at least a dozen individuals from infant to old adult, representation of pretty much the entire skeleton, and a remarkable geological context indicative of intentional disposal of the dead (but certainly not homicide, grumble grumble grumble…). The end of 2015 saw the announcement and uproar (often quite sexist) over this amazing sample. You can expect to see more, positive things about this amazing animal in 2016.

We’ll be presenting a bunch about Homo naledi at this year’s AAPA meeting in Hotlanta. I for one will be discussing dental development at Dinaledi- here’s a teaser:

As long as we’re talking about the AAPA meetings, my colleague David Pappano and I are organizing a workshop, “Using the R Programming Language for Biological Anthropology.” Details to come!

Lemur graveyard

Homo naledi wasn’t the only miraculously copious primate sample announced in 2015. Early last year scientists also reported the discovery of an “Enormous underwater fossil graveyard,” containing fairly complete remains of probably hundreds of extinct lemurs and other animals. As with Homo naledi, such a large sample will reveal lots of critical information about the biology of these extinct species.

Australopithecus deyiremeda

Extended Figure 1h from the paper, with a Demirjian developmental stages, modifed from Table 2 from Kuykendall et al., 1996. Compare the M2 roots with completed roots of the M1 (to the left).

Extended Figure 1h from Haile-Selassie et al. (2015), compared with Demirjian developmental stages 6-8 . While the M1 roots look like stage 8 (complete), M2 looks like stage 7 (incomplete).

We also got a new species of australopithecus last year. Australopithecus deyiremeda had fat mandibles, a relatively short face (possibly…), and smaller teeth than in contemporaneous A. afarensis. One tantalizing thing about this discovery is that we may finally be able to put a face to the mysterious foot from Burtele, since these fossils come from nearby sites of about the same geological age. Also intriguing is the possible evidence, based on published CT images (above), that A. deyiremeda had relatively advanced canine and delayed molar development, a pattern generally attributed to Homo and not other australopithecines (if this turns out to be the case, you heard it here first!).

Lomekwian stone tool industry

3D scan and geographical location of Lomekwian tools. From africanfossils.org.

Roughly contemporaneous with A. deyiremeda, Harmand et al. (2015) report the earliest known stone tools from the 3.3 million year old site of Lomekwi 3 in Kenya. These tools are a bit cruder and much older than the erstwhile oldest tools, the Oldowan from 2.6 million years ago. These Lomekwian tools, and possible evidence for animal butchery at the 3.4 million year old Dikika site in Ethiopia (McPherron et al. 2010; Thompson et al. 2015), point to an earlier origin of lithic technology. Fossils attributed to Kenyanthropus platyops are also found at other sites at Lomekwi. With hints at hominin diversity but no direct associations between fossils and tools at this time, a lingering question is who exactly was making and using the first stone tools.

Earliest Homo

The reconstructed Ledi Geraru mandible (top left), compared with Homo naledi (top right), Australopithecus deyiremeda (bottom left), and the Uraha early Homo mandible from Malawi (bottom right).

The reconstructed Ledi Geraru mandible (top left), compared with Homo naledi (top right), A. deyiremeda (bottom left), and the Uraha early Homo mandible from Malawi (bottom right). Jaws are scaled to roughly the same length from the front to back teeth; the Uraha mandible does not have an erupted third molar whereas the others do and are fully adult.

Just as Sonia Harmand and colleagues pushed back the origins of technology, Brian Villmoare et al. pushed back the origins of the genus Homo, with a 2.7 million year old mandible from Ledi Geraru in Ethiopia. This fossil is only a few hundred thousand years younger than Australopithecus afarensis fossils from the nearby site of Hadar. But the overall anatomy of the Ledi Geraru jaw is quite distinct from A. afarensis, and is much more similar to later Homo fossils (see image above). Hopefully 2016 will reveal other parts of the skeleton of whatever species this jaw belongs to, which will be critical in helping explain how and why our ancestors diverged from the australopithecines. (note that we don’t yet have a date for Homo naledi – maybe these will turn out to be older?)

Early and later Homo

Modified figures X from Maddux et al. (2015) and 13 from Ward et al. (2015).

Left: modified figures 2-3 from Maddux et al. (2015). Right: modified figures 7 & 13 from Ward et al. (2015). Note that in the right plot, ER 5881 femur head diameter is smaller than all other Homo except BSN 49/P27.

The earlier hominin fossil record wasn’t the only part to be shaken up. A small molar (KNM-ER 51261) and a set of associated hip bones (KNM-ER 5881) extended the lower range of size variation in Middle and Early (respectively) Pleistocene Homo. It remains to be seen whether this is due to intraspecific variation, for example sex differences, or taxonomic diversity; my money would be on the former.

Left: Penghu hemi-mandible (Chang et al. 2015: Fig. 3), viewed from the outside (top) and inside (bottom). Right: Manot 1 partial cranium (Hershkovitz et al. 2015: Fig. 2), viewed from the left (top) and back (bottom).

Left: Penghu 1 hemi-mandible (Chang et al. 2015: Fig. 3), viewed from the outside (top) and inside (bottom). Right: Manot 1 partial cranium (Hershkovitz et al. 2015: Fig. 2), viewed from the left (top) and back (bottom).

At the later end of the fossil human spectrum, researchers also announced an archaic looking mandible dredged up from the Taiwan Straits, and a more modern-looking brain case from Israel. The Penghu 1 mandible is likely under 200,000 years old, and suggests a late survival of archaic-looking humans in East Asia. Maybe this is a fossil Denisovan, who knows? What other human fossils are waiting to be discovered from murky depths?

The Manot 1 calvaria looks very similar to Upper Paleolithic European remains, but is about 20,000 years older. At the ESHE meetings, Israel Hershkovitz actually said the brain case compares well with the Shanidar Neandertals. So wait, is it modern or archaic? As is usually the case, with more fossils come more questions.

Crazy dinosaurs

Yi qi was bringing Skeksi back, and its upper limb had a wing-like shape not seen in any other dinosaur, bird or pterosaur. There were a number of other interesting non-human fossil announcements in 2015 (see here and here), proving yet again that evolution is far more creative than your favorite monster movie makers.

ResearchBlogging.org What a year – new species, new tool industries, new ranges of variation! 2015 was a great year to be a paleoanthropologist, and I’ll bet 2016 has just as much excitement in store.

References (in order of appearance)

Haile-Selassie, Y., Gibert, L., Melillo, S., Ryan, T., Alene, M., Deino, A., Levin, N., Scott, G., & Saylor, B. (2015). New species from Ethiopia further expands Middle Pliocene hominin diversity Nature, 521 (7553), 483-488 DOI: 10.1038/nature14448

Harmand, S., Lewis, J., Feibel, C., Lepre, C., Prat, S., Lenoble, A., Boës, X., Quinn, R., Brenet, M., Arroyo, A., Taylor, N., Clément, S., Daver, G., Brugal, J., Leakey, L., Mortlock, R., Wright, J., Lokorodi, S., Kirwa, C., Kent, D., & Roche, H. (2015). 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature, 521 (7552), 310-315. DOI: 10.1038/nature14464

McPherron, S., Alemseged, Z., Marean, C., Wynn, J., Reed, D., Geraads, D., Bobe, R., & Béarat, H. (2010). Evidence for stone-tool-assisted consumption of animal tissues before 3.39 million years ago at Dikika, Ethiopia. Nature, 466 (7308), 857-860. DOI: 10.1038/nature09248

Thompson, J., McPherron, S., Bobe, R., Reed, D., Barr, W., Wynn, J., Marean, C., Geraads, D., & Alemseged, Z. (2015). Taphonomy of fossils from the hominin-bearing deposits at Dikika, Ethiopia Journal of Human Evolution, 86, 112-135 DOI: 10.1016/j.jhevol.2015.06.013

Villmoare, B., Kimbel, W., Seyoum, C., Campisano, C., DiMaggio, E., Rowan, J., Braun, D., Arrowsmith, J., & Reed, K. (2015). Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia Science, 347 (6228), 1352-1355 DOI: 10.1126/science.aaa1343

Maddux, S., Ward, C., Brown, F., Plavcan, J., & Manthi, F. (2015). A 750,000 year old hominin molar from the site of Nadung’a, West Turkana, Kenya Journal of Human Evolution, 80, 179-183 DOI: 10.1016/j.jhevol.2014.11.004

Ward, C., Feibel, C., Hammond, A., Leakey, L., Moffett, E., Plavcan, J., Skinner, M., Spoor, F., & Leakey, M. (2015). Associated ilium and femur from Koobi Fora, Kenya, and postcranial diversity in early Homo Journal of Human Evolution, 81, 48-67 DOI: 10.1016/j.jhevol.2015.01.005

Chang, C., Kaifu, Y., Takai, M., Kono, R., Grün, R., Matsu’ura, S., Kinsley, L., & Lin, L. (2015). The first archaic Homo from Taiwan Nature Communications, 6 DOI: 10.1038/ncomms7037

Hershkovitz, I., Marder, O., Ayalon, A., Bar-Matthews, M., Yasur, G., Boaretto, E., Caracuta, V., Alex, B., Frumkin, A., Goder-Goldberger, M., Gunz, P., Holloway, R., Latimer, B., Lavi, R., Matthews, A., Slon, V., Mayer, D., Berna, F., Bar-Oz, G., Yeshurun, R., May, H., Hans, M., Weber, G., & Barzilai, O. (2015). Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans Nature, 520 (7546), 216-219 DOI: 10.1038/nature14134

Bioanthro lab activity: What species is it?

October 30, 2015October 30, 2015 / zcofran / Leave a comment

We’re learning about the divergence between robust Australopithecus and early Homo 2.5-ish million years ago in my Human Evolution class this week. Because of this multiplicity of contemporaneous species, when scientists find new hominin fossils in Early Pleistocene sites, a preliminary question is, “What species is it?”

Scrutinizing the fossil record, asking the difficult questions. (Science credit)

To help my students learn how we know whether certain fossils belong to the same species, and to which group new fossils might belong, in this week’s lab we compared tooth sizes of Australopithecus boisei and early Homo. After seeing how tooth sizes differed between these groups, students then tested whether they could determine whether two “mystery” fossils (KNM-ER 60000 and 62000; Leakey et al. 2012) belonged either group.

Early Pleistocene hominin fossils from Kenya. Left to right: KNM-ER 406, ER 62000 and ER 1470. At the center is one f the lab’s “mystery jaws.”

Students downloaded 3D scans of hominin fossils from AfricanFossils.org, and measured buccolingual/labiolingual tooth crown diameters using MeshLab.

Early Pleistocene hominin mandibles. Left to right: KNM-ER 3230, ER 60000 (“mystery” jaw) and ER 1802.

The first purpose of this lab was to help familiarize students with skull and tooth anatomy of early Pleistocene humans. Although lectures and readings are full of images, a lab activity forces students to spend time visually examining fossils. Plus, they’re in 3D which is a whole D greater than 2D – the visual equivalent of going to eleven! The second goal of the lab was to help prepare students for their term projects, in which they must pose a research question about human evolution, generate predictions, and find and use data to test hypotheses.

If you’re interested in using or adapting this activity for your class, here are the handout and data sheet into which students enter their measurements. The data sheet specifies the fossils that can be downloaded from africanfossils.org. Some relevant fossils (i.e., KNM WT 15000 and ER 992) were not included because the 3D scans yield larger measurements than in reality.

Lab 3-Mystery Jaws (instructions and questions)

Lab 3-Mystery jaws data sheet

Reference
Leakey MG, Spoor F, Dean MC, Feibel CS, Antón SC, Kiarie C, & Leakey LN (2012). New fossils from Koobi Fora in northern Kenya confirm taxonomic diversity in early Homo. Nature, 488 (7410), 201-4 PMID: 22874966

eFfing #FossilFriday: Rekindling an old friend’s hip

June 12, 2015January 1, 2016 / zcofran / 2 Comments

Sorry for the crappy pun. Carol Ward and colleagues recently reported an associated hip joint, KNM-ER 5881, attributable to the genus Homo (1.9 million years old). Fossils coming from the same skeleton are pretty rare, but what’s more remarkable is that portions of this bone were discovered 29 years apart: a femur fragment was first found in 1980, and more of the femur and part of the ilium were found at the same location when scientists returned in 2009:

Figure 3 from Ward et al. 2015. A little distal to the hip, yes, but the pun still works. Views are, going clockwise starting at the top the top left, from above, from below, from the back, from the side, and from the front.

There’s also a partial ilium associated with the femur – that makes a pretty complete hip!

Figure 5 from Ward et al. shows the fossil. Jump for joy that it’s complete enough for us to tell it comes from the left side!

Despite how fragmentary the femur and ilium are, the researchers were able to estimate the diameter of the femur head and hip socket reliably. The hip joints are smaller than all Early Pleistocene Homo except for the Gona pelvis. Comparing ER 5881 the large contemporaneous KNM-ER 3228 hip bone, the authors found these two specimens to be more different in size than is usually seen between sexes of many primate species. The size difference best matches male-female differences in highly dimorphic species like gorillas.

Ward et al. find that the specimen generally looks like early Homo but that the inferred shape of the pelvic inlet is a little different from all other Early and Middle Pleistocene human fossils. The authors take this discrepancy to suggest that there was more than one “morphotype” (‘kind of shape’), and therefore possibly species, of Homo around 1.9 million years ago. While I wouldn’t just yet go so far as to say this anatomy is due to species differences, I do agree that KNM ER 5881 helps our understanding and appreciation of anatomical variation in our early ancestors. Like all great fossil discoveries, the more we find, the more we learn that we don’t know. Here’s to more Homo hips in the near future!

eFfing #FossilFriday: Pleistocene ppl blowin up this week

January 30, 2015February 22, 2017 / zcofran / 2 Comments

This was a big week for Middle-Late Pleistocene fossil humans. Chun-Hsiang Chang and colleagues describe a mandible dredged up off the western coast of Taiwan, which they note in the title as, “The first archaic Homo” fossil known from the region. The geological context makes it difficult to date the specimen precisely, but authors argue it is probably younger than 190 thousand years old.

The Penghu mandible. Figure 3. From Chang et al.

In life, this individual was fully grown but appears never to have developed third molars (the “wisdom teeth”). Such “third molar agenesis” is relatively rare before modern times, but is also seen in the D2735 Homo erectus mandible from Dmanisi. I wouldn’t make much of this coincidence, but it does raise the question of whether the cause of agenesis, not uncommon today, was the same then as now.

Shortly after the announcement of the Penghu mandible, Israel Hershkovitz and colleagues presented a 55,000 year old brain case from Manot Cave in the Levant. The calvaria (fancy word for brain case) looks very similar to the skulls of the slightly younger “anatomically modern” humans of the Upper Paleolithic in Europe, albeit with a few Neandertal-like traits here and there (hey, just like many of the Upper Paleolithic humans).

The Manot calvaria (Figure 2 from Hershkovitz et al.) The views are (a-d) from the top with front to the left; from the left; from the front; and from the back. Extra credit: In the top view (a), can you identify the features telling that the front is to the left?

John Hawks has good posts dedicated to both Penghu and Manot. The upshot of these discoveries is that Middle and Late Pleistocene human population diversity, and the interactions between these populations, are probably much more complicated and interesting than the old model of ‘modern’ humans arising singly in Africa and replacing ‘archaic’ humans in different parts of the globe. With the technological advances and fossil discoveries of the past decade, the rather simple Replacement model has given way to a better appreciation of true complexity of human evolution toward the end of the Ice Age. Both of these new papers reflect this new perspective.

Along these lines, accompanying the Manot paper in Nature is an editorial, “Human history defies easy stories.” What caught my attention reading this (anonymous?) commentary is that it puts scientific interpretations of the past into a social and historical context. The author notes that the traditional story of modern humans arising, spreading and eradicating other groups of human has “imperialist framing, in which evolution and replacement can be justified after the fact as a kind of manifest destiny.” Science doesn’t occur in a vacuum, it’s done by people whose minds and creativities are molded in specific historical, economic and cultural contexts. This editorial comment makes one wonder how the human fossil record would have been interpreted, had most of it not discovered against the social backdrop of ruthless capitalism.

eFfing Fossil Friday: resurrected

June 20, 2014June 20, 2014 / zcofran / 2 Comments

It’s been a quiet month here at Lawnchair, as I’ve just returned from the Rising Star Workshop, taking part in the analysis and description of new hominin remains from South Africa. We’ll have some exciting announcements to make in the near future.

Also, I petted a ferocious, bloodthirsty lion!

To ease back into the Lawnchair, I thought I’d resurrect eFfing Fossil Friday, a short-lived series from when I was collecting data for my dissertation three years ago (speaking of which, a paper related to my dissertation came out in AJPA during the Workshop, as well). A lot has happened since the last installment of FFF, so whose heads will be on the chopping block today?

Crania 9, 15 and 17 (clockwise from top left). Cranium 9 is an early adolescent and the other two are adults - lookit how the facial anatomy changes with age!

Crania #s 9, 15 and 17 (clockwise from top left). Cranium 9 is an early adolescent and the other two are adults – lookit how the facial anatomy changes with age! (Fig. 1 from Arsuaga et al., 2014)

It’s new crania from Sima de los Huesos, Atapuerca! These are published today in the journal Science by Juan L. Arsuaga and colleagues. Sima de los Huesos is a pretty remarkable site in Spain dating to the Middle Pleistocene; the site is probably at least 400,000 years old, and the remains of at least 28 individuals. These specimens show many similarities with Neandertals who later inhabited the area, but don’t have all of the ‘classic’ Neandertal features.

What I like about this figure from the paper is that the comparison of the adolescent (top left) with adults (the other two) shows how the skull changes during growth. The major visible difference is that the face sticks out in front of the brain case more in the adults than the adolescent. As a result, the adolescent lacks a supraorbital torus (“brow ridge”), but this would have developed as the face grew forward and away from the brain. Ontogeny!

Gona … Gona … not Gona work here anymore more

March 4, 2014March 5, 2014 / zcofran / 2 Comments

The Gona pelvic remains (A-D), and the reconstructed complete pelvis (E-J), Fig. 2 in Simpson et al., 2008.

A few years ago, Scott Simpson and colleagues published some of the most complete fossil human hips (right). The fossils are from the Busidima geological formation in the Gona region of Ethiopia, dated to between 0.9-1.4 million years ago. (Back when I wasn’t the only author of this blog, my friend and colleague Caroline VanSickle wrote about it here)

Researchers attributed the pelvis to Homo erectus on the basis of its late geological age and a number of derived (Homo-like) features. In addition, the pelvis’s very small size indicated it probably belonged to a female. One implication of this fossil was that male and female H. erectus differed drastically in body size.

Christopher Ruff (2010) took issue with how small this specimen was, noting that its overall size is more similar to the small-bodied Australopithecus species. Using the size of the hip joint as a proxy for body mass, Ruff argued Gona’s small size would imply a profound amount of sexual dimorphism in H. erectus: much higher than if Gona is excluded from this species, and higher than in modern humans or other fossil humans. Ruff thus proposed an alternative hypothesis to marked sexual dimorphism, that the Gona pelvis may have belonged to an australopithecine.

Fig. 3 From Ruff's (2010) reply. Australopiths (and Orrorin) are squares and Homo are circles. Busidima's estimated femur head diameter is represented by the star and bar.

Fig. 3 From Ruff’s (2010) reply. Australopiths (and Orrorin) are squares and Homo are circles. Gona’s estimated femur head diameter is represented by the star and bar.

Now, Simpson & team replied to Ruff’s comments, providing a laundry list of reasons why this pelvis is H. erectus and not Australopithecus. They cite many anatomical features of the pelvis shared with Gona and Homo fossils, but not australopithecines. They also note that there are many other bones reflective of body size, that seem to suggest a substantial amount of size variation in Homo fossils, even those from a single site such as Dmanisi (Lordkipanadze et al., 2007).

Interestingly, neither of these parties compared the implied size variation with that of living apes. So I’ll do it! Now, I do not have any acetabulum data, but a friend lent me some femur head measurements for living great apes a few years ago. Gona is a pelvis and not a femur, but there are more fossil femora than hips. Because there’s a very high correlation between femur head and acetabulum size, Ruff estimated Gona’s femur head diameter to be 32.6 mm (95% confidence interval: 30.1-35.2; Simpson et al. initially estimated 35.1 mm based on a different dataset and method). To quantify size variation, we can compare ratios of larger femur heads divided by smaller ones. Now, this ratio quantifies inter-individual variation, but it will underestimate sexual dimorphism since I’m likely sampling some same-sex pairs that aren’t so different in size. But this is just a quick and dirty look. So, here’s a box plot of these ratios for Homo fossils, larger specimens divided by Gona’s estimated femur head size in different time periods:

Ratio of a fossil Homo femur head diameter (HD) divided by Busidima's HD. E Homo = early Pleistocene, Contemporaneous = WT 15000 and OH 28, MP = Middle Pleistocene Homo. White boxes are based on Ruff's Busidima HD estimate, green boxes are based on Simpson et al.'s estimate.

Ratios of fossil Homo femur head diameter (HD) divided by Busidima’s (Gona’s) HD. E Homo = early Pleistocene, Contemporaneous = WT 15000 and OH 28, MP = Middle Pleistocene Homo. White boxes are based on Ruff’s Gona HD estimate, green boxes are based on Simpson et al.’s larger estimate. Boxes include 50% quartiles and the thick lines within are sample medians.

Clearly, Gona is much smaller than most other fossil Homo hips, since ratios are never smaller than 1.14. Average body size increases over time in the Homo lineage, reflected in increasing ratios from left to right on the plot. Early Pleistocene Homo fossils are fairly small, including Dmanisi, hence the lower ratios than later time periods. Middle Pleistocene Homo (MP), represented by the most fossils, shows a large range of variation, but even the smallest is still 1.17 times larger than the largest estimate of Gona’s femur head size. To put this into context, here are those green ratios (assuming a larger size for Gona) compared with large/small ratios from resampled pairs of living apes and humans:

The fossil ratios of larger/smaller HD from above, compared with resampled ratios from unsexed living apes and humans. Boxes include the 50% quartiles, and the thick lines within are sample medians. **(05/03/14: This plot has been modified from the original version post, which only included the fossil ratios based on the smaller Gona estimate)

What we see for the extant apes and humans makes sense: humans and chimpanzees show smaller differences on average, whereas average differences between gorillas and orangutans are larger. This accords with patterns of sexual dimorphism in these species. **What this larger box plot shows is that if we accept Ruff’s smaller average estimate of Gona’s femur head size (white boxes), it is relatively rare to sample two living specimens so different in size as seen between Gona and other fossils. If we use Simpson et al.’s larger Gona size estimate, variation is still elevated above most living ape ratios. Only when Gona is compared with the generally-smaller, earlier Pleistocene fossils, does the estimated range of variation show decent overlap with living species. Even then, the overlap is still above the median values.

These results based on living species agree with Ruff’s concern, that including Gona in Homo erectus results in an unusually large range of variation in this species. Such a large size range isn’t necessarily impossible, but it would be surprising to see more variation than is common in gorillas and orangutans, where sexual size dimorphism is tremendous. Ruff suggested that the australopith-sized Gona pelvis may in fact be an australopith. This was initially deemed unlikely, in part because the fossil is well-dated to relatively late, 0.9-1.4 million years ago. However, Dominguez-Rodgrigo and colleauges (2013) recently reported a 1.34 mya Australopithecus boisei skeleton from Olduvai Gorge, so it is possible that australopiths persisted longer than we’ve got fossil evidence for, and Gona is one of the latest holdouts.

So many possible explanations. More clarity may come with further study of the fossils at hand, but chances are we won’t be able to eliminate any of these possibilities until we get more fossils. (also, the post title wasn’t a jab at the fossils or researchers, but rather a reference to the movie Office Space)

References

Dominguez-Rodrigo et al. 2013. First partial skeleton of a 1.33-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania. PLoS One 8: e80347.

Ruff C. 2010. Body size and body shape in early hominins – implications of the Gona pelvis. Journal of Human Evolution 58: 166-178.

Simpson S et al. 2008. A female Homo erectus pelvis from Gona, Ethiopia. Science 322: 1089-1092.

Simpson S et al. In press. The female Homo pelvis from Gona: Response to Ruff (2010). Journal of Human Evolution. http://dx.doi.org/10.1016/j.jhevol.2013.12.004

Evo-devo of the human shoulder?

January 1, 2012August 4, 2013 / zcofran / 1 Comment

It’s a new year, and while my mind should be marred by a hangover, instead all I can think about are fossils and scapulas.

A pretty cool study was published online in the Journal of Human Evolution last week, and I’ve finally gotten to peruse it. Fabio Di Vincenzo and colleagues analyzed the shape of the outline of the glenoid fossa on the scapula (not to be confused with the glenoid on your skull), from Australopithecus africanus to present day humans. The glenoid fossa is essentially the socket in the ball-and-socket joint of your shoulder. The authors found that there is pretty much a single trend of glenoid shape change from Australopithecus through the evolution of the genus Homo: from the fairly narrow joint in Australopithecus africanus and A. sediba, to the relatively wide joint in recent humans. The overall size and shape of the joint influences/reflects shoulder mobility, so presumably this shape change hints that more front-to-back arm motions became more important through the course of human evolution (authors suggest throwing in humans from the Late Pleistocene onward).

The finding of a single predominant trend in glenoid shape evolution is pretty interesting. On top of that, the authors add an ‘evo-devo’ twist by comparing species’ average “shapes” (first principle component scores, on the y-axis in the figure at right) with their estimated ages at skeletal maturity (which appears scaled to the modern human value, on the x-axis). Though it’s not an ideal dataset for running a linear regression, the figure at right shows that there appears to be a fairly linear relationship across human evolution, such that groups with an older age at skeletal maturity tend to have a more rounded (modern human-like) glenoid fossa (note that the individuals in the analysis were all adults). Overall size does not contribute to shape variation among these glenoids.

This work raises two issues, and ultimately leads to a testable evo-devo hypothesis. The first issue is to what extent we can trust their estimates of age at skeletal maturity. These estimates were allegedly taken from a chapter by Helmut Hemmer (2007) in the prohibitively expensive Handbook of Paleoanthropology. Cursorily glancing at this chapter, I can’t find age at skeletal maturation estimated for any hominids. It is possible that in my skimming I missed the estimates, or alternatively that Di Vincenzo and colleagues misinterpreted another variable as skeletal development. Either way, these estimates would still need to be taken with a grain of salt, given that it is almost impossible to know the true age at death of a fossil (but see Antoine et al. 2008), especially if there are no associated cranio-dental elements.

That said, it is perfectly reasonable to suppose that the age at skeletal maturation has increased over the course of human evolution; life-span increased through human evolution, and so all else being equal (which it almost certainly isn’t) we could expect that maturation would occur later over time, too. So this leads to a second issue: given the “evo-devo change” the authors hypothesize, what is the evo-devo mechanism? That is, how was development modified to effect the evolutionary changes we see in the hominid scapula? Because they found adult glenoid shape correlates with estimated age at skeletal maturity, this leads to the hypothesis that postnatal skeletal growth accounts for the shape difference. Indeed, they state:

“If functional and static allometric influences are unlikely, we…interpret the trend…as reflecting growth and developmental factors. A major, albeit gradual, trend of ontogenetic heterochrony occurred in the evolution of the genus Homo… and thus differences within and between taxa in overall growth rates may have produced the pattern of variation between samples, as well as the overall temporal trend observed. The regression of life history variables [they only looked at 1]… with PCA [principle components analysis] scores supports this ‘ontogenetic’ hypothesis.”

The authors suggest that humans’ slower growth rates but longer growth period “led to longer periods of bone deposition along the inferior-lateral edge of the [glenoid fossa]” The heterochronic process they suggest is “peramorphosis” – the descendant reaches a shape that is ‘beyond’ that of the ancestor.

The figure above is from a seminal “heterochrony” paper by Pere Alberch and colleagues (1979), portraying how peramorphosis can occur. In each, the y-axis represents shape and the x-axis is age. A the descendant’s peramorphic shape (“Ya”) could result from accelerated growth (left graph) or from an extension of growth to later ages than in the ancestor (right graph).

And so this leads to a testable hypothesis. Di Vincenzo and colleagues cite (dental) evidence that humans’ overall body growth rates are slower than earlier hominids’, undermining the hypothesis that acceleration is responsible for humans’ glenoid peramorphosis. Rather, they hypothesize that humans’ slower growth rates coupled with a longer period of skeletal development, to result in a relatively wider glenoid, due to increased development of the secondary growth centers (e.g. the graph at right, above). This developmental scenario predicts that subadult human glenoids should resemble earlier hominid adults’, that “ontogeny recapitulates phylogeny” as far as glenoid shape is concerned. Analyzing glenoid growth can even be extended to include fossils – the >3 million year old human ancestor Australopithecus afarensis has glenoids preserved for an infant (DIK-VP-1; Alemseged et al. 2006) and 2 adults (AL 288 “Lucy” and KSD-VP-1; Johanson et al. 1982, Haile-Selassie et al. 2010). An alternate hypothesis is that species’ distinct glenoid shapes are established early during life (i.e. in utero), and/or that no simple heterochronic process is involved.

Di Vincenzo’s and colleagues’ study points to the importance of development in understanding human evolution, and their hypothesized “evo-devo change” in glenoid shape is ripe for testing.

References
Pere Alberch, Stephen Jay Gould, George F. Oster, & David B. Wake (1979). Size and shape in ontogeny and phylogeny Paleobiology, 5 (3), 296-317

Alemseged, Z., Spoor, F., Kimbel, W., Bobe, R., Geraads, D., Reed, D., & Wynn, J. (2006). A juvenile early hominin skeleton from Dikika, Ethiopia Nature, 443 (7109), 296-301 DOI: 10.1038/nature05047

Antoine, D., Hillson, S., & Dean, M. (2009). The developmental clock of dental enamel: a test for the periodicity of prism cross-striations in modern humans and an evaluation of the most likely sources of error in histological studies of this kind Journal of Anatomy, 214 (1), 45-55 DOI: 10.1111/j.1469-7580.2008.01010.x

Di Vincenzo, F., Churchill, S., & Manzi, G. (2011). The Vindija Neanderthal scapular glenoid fossa: Comparative shape analysis suggests evo-devo changes among Neanderthals Journal of Human Evolution DOI: 10.1016/j.jhevol.2011.11.010

Haile-Selassie, Y., Latimer, B., Alene, M., Deino, A., Gibert, L., Melillo, S., Saylor, B., Scott, G., & Lovejoy, C. (2010). An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia Proceedings of the National Academy of Sciences, 107 (27), 12121-12126 DOI: 10.1073/pnas.1004527107

Hemmer, Helmut (2007). Estimation of Basic Life History Data of Fossil Hominoids Handbook of Paleoanthropology, 587-619 DOI: 10.1007/978-3-540-33761-4_19

Johanson, D., Lovejoy, C., Kimbel, W., White, T., Ward, S., Bush, M., Latimer, B., & Coppens, Y. (1982). Morphology of the Pliocene partial hominid skeleton (A.L. 288-1) from the Hadar formation, Ethiopia American Journal of Physical Anthropology, 57, 403-451 DOI: 10.1002/ajpa.1330570403

[insert clever quip about australopithecus hips]

September 19, 2011August 4, 2013 / zcofran / Leave a comment

A week and a half ago, Kibii and colleagues (2011) published reconstructions and re-analyses of two hips belonging to the 1.98 million-year old Australopithecus sediba. As with many fossil discoveries, these additions to the fossil record raise more questions than they answer. Unless the question was, “did A. sediba have a pelvis?” It did. Here’s a good summary from the paper itself:

Thus, Au. sediba is australopith-like in having a long superior pubic ramus and an anteriorly positioned and indistinctly developed iliac pillar…[and] Homo-like in having vertically oriented and sigmoid shaped iliac blades, more robust ilia, and a narrow tuberoacetabular sulcus…and the pubic body is upwardly rotated as in Homo. (p. 1410, emphases mine)

So far as I can tell, the main way the hips are ‘advanced’ toward a more human-like condition is that the iliac blades are more upright and sweep forward more than in earlier known hominid hips. Here’s the figure 2 from the paper (more sweet pics of the fossils are available here). NB that in both A. sediba hips much of the upper portions of the iliac blades are missing (reconstructed in white; this region is missing in lots of fossils), so it’s possible they were more flaring like the australopith in the center photo.

The authors’ bottom-line, take-home point is that the A. sediba pelvis has features traditionally associated with large-brained Homo – but belonged to a small-brained species (based solely on the ~430 cc MH1 endocast). They argue that this means that many of these unique pelvic features did not evolve in the context of birthing large-brained babies, as has often been thought. They state that these features are thus “most parsimoniously attributed to altered biomechanical demands on the pelvis in locomotion,” and suggest that this hypothetical locomotion was mostly bipedalism but with a good degree of climbing. Maybe, maybe not. This interpretation is consistent with the analysis of the A. sediba foot/ankle (Zipfel et al. 2011).

The weird mix of ancient (australopith-like) and newer (Homo-like) pelvic features in A. sediba really raises the question of how australopithecines moved around. More intriguing is that the A. sediba pelvis has different Homo-like features than the ~1 million year old Busidima pelvis (Simpson et al. 2008), which has been attributed to Homo erectus (largely in aspects of the iliac blades). This raises the question of whether A. sediba is really pertinent to the origins of the genus Homo, and whether the Busidima pelvis belongs to Homo erectus or a late-surviving robust australopithecus (e.g. boisei, Ruff 2010).

Also interesting is that the subpubic angle (in the pic above, the upside-down “V” created by the pubic bones just above the red labels) is pretty low in MH2. This is curious because modern human males and females differ in how large this angle is – females tend to have a large angle which contributes to an enlarged birth canal, whereas males have a low angle like MH2. But MH2 is considered female based on skeletal and dental size. This raises the additional questions of whether human-like sexual dimorphism had not evolved in hominids prior to 1.9 million years ago, and whether the sex of MH2 was accurately described.

Finally, though the authors did a great job comparing this pelvis with those from other hominids, I think a major, more comprehensive comparative review of hominid pelves is in order. How does the older A. afarensis hip from Woranso (Haile-Selassie et al. 2010) inform australopithecine pelvic evolution? What about the possibly-contemporary-maybe-later hip from the nearby site of Drimolen (Gommery et al. 2002)? Given the subadult status of the MH1 individual, it would be interesting to compare with the WT 15000 Homo erectus fossils, or A. africanus subadults from Makapansgat, to examine the evolution of pelvic growth.

Lots of interesting questions arise from these fascinating new fossils. “The more you know,” right?

References
Gommery, D. (2002). Description d’un bassin fragmentaire de Paranthropus robustus du site Plio-Pléistocène de Drimolen (Afrique du Sud)A fragmentary pelvis of Paranthropus robustus of the Plio-Pleistocene site of Drimolen (Republic of South Africa) Geobios, 35 (2), 265-281 DOI: 10.1016/S0016-6995(02)00022-0

Haile-Selassie Y, Latimer BM, Alene M, Deino AL, Gibert L, Melillo SM, Saylor BZ, Scott GR, & Lovejoy CO (2010). An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia. Proceedings of the National Academy of Sciences of the United States of America, 107 (27), 12121-6 PMID: 20566837

Kibii, J., Churchill, S., Schmid, P., Carlson, K., Reed, N., de Ruiter, D., & Berger, L. (2011). A Partial Pelvis of Australopithecus sediba Science, 333 (6048), 1407-1411 DOI: 10.1126/science.1202521

Ruff, C. (2010). Body size and body shape in early hominins – implications of the Gona Pelvis Journal of Human Evolution, 58 (2), 166-178 DOI: 10.1016/j.jhevol.2009.10.003

Simpson, S., Quade, J., Levin, N., Butler, R., Dupont-Nivet, G., Everett, M., & Semaw, S. (2008). A Female Homo erectus Pelvis from Gona, Ethiopia Science, 322 (5904), 1089-1092 DOI: 10.1126/science.1163592

Zipfel, B., DeSilva, J., Kidd, R., Carlson, K., Churchill, S., & Berger, L. (2011). The Foot and Ankle of Australopithecus sediba Science, 333 (6048), 1417-1420 DOI: 10.1126/science.1202703

100,000 year old child skeleton on National Geographic

June 17, 2011June 9, 2023 / zcofran / Leave a comment

National Geographic aired a special tonight about a recently-excavated child’s skeleton (they focused on the skull) from Grotte des Contrebandiers in Morocco, dated to around 108,000 years ago. So far as I know this material has not been fully published (aside from a brief blurb in Science).

The program presented work of archaeologists, paleontologists, reconstruction artists, taphonomists, and lots of other people, hoping to figure out who the kid was and such. All in all it was pretty cool, I’d recommend checking it out if you didn’t see it. Or again if you did see it.

While I think it was a great program and the researchers involved are doing a terrific job, I had two main concerns: first, I wish they’d treated the topic of growth-n-development a little more. They noted that the child (5-6 years old possibly) looked really “modern” because of its flat face. But looking at it, it didn’t really have that flat of a face, especially for a child. They talked about how human-like (rather than Neandertal-like) the kid was, but they only compared it with adults – children tend to have relatively smaller faces and larger brain-cases than adults (right), so it’s no wonder it looked more like an adult human than the adult Neandertal from Amud (Israel) that they compared it with. It would’ve been great to see more comparisons with other late Pleistocene hominid kids, such as from Skhul/Qafzeh or La Quina. A future program, perhaps.

Second, they kept asking whether the kid was “a Homo sapien.” I know it’s counterintuitive for English-speakers, but “H. sapiens” is the singular and plural of humans’ scientific name. Silly, right, cuz it doesn’t even get paid twice as much. But you’ll have take that up with C. Linnaeus. I am a Homo sapiens. You are a Homo sapiens. Fifty people are a gaggle of Homo sapiens.

Anyway it was a cool show. Check it out, dammit!

Figure credit: Fig. 2 from Bogin. 2003. The human pattern of growth and development in paleontological perspective. In Patterns of Growth and Development in the Genus Homo, eds. Thompson JL, Krovitz GE and Nelson AJ. New York: Cambridge University Press: 15-44.

Holy Effing Crap II: Australopithecus from Malapa

April 8, 2010 / zcofran / Leave a comment

Lee Berger and colleagues report in Science today on 2 incredibly well preserved skeletons – including perhaps the best-preserved hominid skull in South Africa, in some ways as good as or better than Sts 5 (Australopithecus africanus). The specimens come from a site called Malapa in South Africa, dating to around 1.9 – 1.7 million years ago. The authors argue that it is so unique in its features that it warrants a new species – Australopithecus sediba – linking the earlier A. africanus with later Homo habilis. Is it really a new species? In my personal opinion, there’s not much distinguishing it from A. africanus.

The amazingly preserved skull is of a subadult, maybe 11 years old. The highly angled root of the zygomatic, positioned just above the M1 alveolus is classic A. africanus. It really reminds me of Sts 17, or possibly Sts 52 in the lower face. The prognathism is modest and lacks the anterior nasal pillars which tend to be fairly pronounced in A. africanus; in this regard it is quite comparable to specimens like TM 1512. Like Sts 52, it has multiple infraorbital foramina. The cranial capacity is estimated at 420 cubic centimeters (cc), which is pretty small, but within the estimated A. africanus range of variation. The authors say that the relatively low position of the temporal lines, spaced far apart from the sagittal suture, and the fact that the zygomatic arches do not flare to the sides, are features more like Homo than like A. africanus. But the specimen is only 11 years old; while the brain is finished growing, the face and chewing muscles probably aren’t. So if this were a fully adult specimen, I’m sure both of these features would come to look more like A. africanus.

The upper and lower first and second molars increase in size posteriorly, and have a distinct protostylid (enamel shelf along the side of one of the cusps) which has a very high frequency in A. africanus. The upper molars, while not totally complete, preserve something that I’ve noticed and I’m sure is in the literature, that the M1 is fairly small and squared compared to the generally larger and not-quite-as-square M2. In a few words, then, the skull seems to fit comfortably in the range of A. africanus variation.

Perhaps the least A. africanus-like aspect of the skull is the supraorbital torus. The supraorbital, or brow, is generally a modestly expressed in most africanus specimens that preserve it. The Malapa specimen is much more similar, to my eye, to later Homo in its projection and arching over the eyes. What could this mean? Moss and Young’s (1960) functional matrix model of looking at the cranium views the supraorbital as a function of the relative position of the brain to the orbits. Perhaps the spatial relationship between the vault and the face which becomes characteristic of later homo becomes established in earlier in the lineage. Other than the supraorbital, this specimen seems purely A. africanus to me. In all, the contour of the vault may not be too different from younger Dmanisi specimens like D2700 or 2280; that Malapa lacks the occiput gives an artificially short front-to-back look to the specimen. The face, however, is totally A. africanus.

Perhaps one of the most striking images is the lateral view. This photo looks strikingly similar to a subadult chimpanzee, albeit with a taller face and less prognathic snout. Maybe I’ve just seen a subadult ape before that this thing reminds me of.

So, this is an immensely exciting set of fossils. Is there a new species of Australopithecus? I wouldn’t bet my life on it. If you go with the widely held idea, that A. africanus or something like it was ancestral to later A. robustus on the one hand, and our Homo ancestors on the other, this thing would fall on the Homo side of that split. So in this case, since we’re not seeing anagenetic evolution – evolutionary change within a lineage – but rather branching, how do you name this thing? It might be slightly more derived toward a Homo than either its “pure” africanus ancestors and A. robustus evolutionary cousins, does this make it Homo? The issue is that adaptively its morphology doesn’t seem to be different from A. africanus, which would argue against the generic distinction. But if its later ancestors become H. habilis and nothing else, then I suppose this would make it a “chronospecies” of H. habilis. So maybe we should call this thing H. habilis? I think most people would argue with this simply on the brain size issue. And the brain is way smaller than any proclaimed Homo specimen.

Taxonomically, this will be a tough call.