#FossilFriday: 2015 Retrospecticus


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

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

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

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.orgWhat 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?

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?”

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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.

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.

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

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

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.

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!

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

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?

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 #FossilFriday: Subfossil lemurs

Hard to resist the headline, “Enormous underwater fossil graveyard found,” from the National Science Foundation. The NSF posts a video detailing the discovery of an underwater cave system containing “hundreds of potentially 1,000-year-old [lemur] skeletons…” in Madagascar. As a paleontologist, hearing about the discovery large numbers of ancient skeletons is musical, like hearing Love This Giant or the new T Swift for the first time.

Two lemur crania in an underwater cave on Madagascar. Photo from nbcnews.com.

Two lemur crania in an underwater cave on Madagascar. Photo from nbcnews.com.

It’s a pretty remarkable discovery – hundreds if not thousands of bones representing many complete skeletons of various extinct lemur species. And toward the end of the clip is a skull of a pretty badass looking big cat. The video shows piles of loose bones dredged up from the cave. These will reveal lots of information about the biology of these recently extinct animals, especially if researchers can keep associated bones together.

So what are these animals? Lemurs are one of the most primitive living types of primates – although they are relatively closely related to us humans, they retain many characteristics of ancestral mammals. I know it’s hard to believe this aye-aye here is more closely related to you than to rodents, but it is:

An aye-aye (Daubentonia madagascarensis) using its narrow and elongated middle finger to fish for for grubs inside a tree that it’s opened up with its teeth.

Lemurs are found only on the island of Madagascar, and over the past several millions of years they have diversified into the roughly 100 species inhabiting the island today. But even just a few thousand years ago, there were more kinds of lemurs. This includes Megaladapis, the large-bodied “koala lemur,” and Hadropithecus, whose skull bears a striking resemblance to the extinct hominin Australopithecus boisei. As  Laurie Godfrey says in the video, “two thirds of the animals that lived there only a thousand years ago are gone.” Humans are probably largely responsible for the extinction of many Malagasy lemurs in both the past and especially the present.

Much of the ‘fossil’ record for lemurs is recent by fossil standards, and so most specimens haven’t become fully fossilized. As a result, lemur paleontology is besprinkled with the term “subfossil,” indicating bones that are really old and belong to extinct animals, but don’t fit the technical definition of fossils. The lemur subfossil record has taught us a lot about the evolutionary history, adaptations, and recently even genetics of this primitive group of primates, as well as about the ecological history of Madagascar. It will be very interesting to see what new insights will come from the recently discovered scores of underwater skeletons.


(Figure 3 from Ryan et al., 2008. Scale bar is 1 cm)

eFfing Fossil Friday: Funky facial flanges #FFF

David Krause and colleagues announced in this week’s Nature the discovery of a new species of extinct mammal, Vintana sertichi, that lived in what is now Madagascar between 66-72 million years ago. The species is based on a very well-preserved cranium of an early gondwanatherian (if you want to impress your friends this weekend, gratuitously use the word “gondwanatherian”). I don’t know much about early mammals like this, but it sounds like it was a weird creature (see the Stony Brook press release). Just looking at it’s face there’s something that sticks out as strange:

Ventana sertichi cranium (Reich et al. 2014, Figure 1a). Left is a 3D CT reconstruction, right is a line drawing highlighting all the individual bones (so many cranial bones). The view is from the right side, so the nose is on the right, the eye is the big hollow in the middle, and the back of the skull is on the left. The jugal flanges are the downward projections.

Vintana sertichi cranium (Reich et al. 2014, Figure 1a). On the left is a 3D CT reconstruction, and on the right is a line drawing highlighting all the individual bones (so many cranial bones). The view is from the right side, so the nose is to the right, the eye socket is the shadowy hollow in the middle, and the back of the skull is on the left. The jugal flanges are the downward projections.

Jutting downward from the sides of the jaw are ‘jugal flanges,’ projections of bone on the homologs of human cheeks. Projections of like these usually serve as muscle attachment sites, and the size of the projection generally reflects the size of the muscle. These facial flanges anchor the masseter muscle, a major chewing muscle that helps close the jaw. The size of this flange in Vintana suggests its chomp packed a punch. A debilitating bite. A face not even a mother could love (so now they’re extinct).

Vintana‘s bony tear-catchers caught my eye because most primates I’ve seen have, you know, less heinous faces. Scouring the internet, big jugal flanges are a fairly rare sight, but can apparently be found in glyptodonts (giant, armadillo-like mammals that lived tens of thousands of years ago) and various sloths. The closest thing I’ve seen to this gross bony flange in Primates are on the zygomatic bones of some extinct, baboon-like animals, such as Dinopithecus ingens:

Fragmentary skull, viewed from the top, of Papio (a.k.a. Dinopithecus) ingens, from Swartkrans, South Africa. Photo credit: CalPhotos.

Fragmentary skull, viewed from the top, of Papio (a.k.a. Dinopithecus) ingens, from Swartkrans, South Africa. As a punishment for its zygomatic excess, its face was confiscated. Photo credit: CalPhotos.

and Theropithecus brumpti

Theropithecus brumpti from the Omo basin. Photo credit: CalPhotos.

Theropithecus brumpti from the Omo Basin, Ethiopia. Photo credit: CalPhotos.

So some primates dabbled in jugal flangery like Vintana, but Natural Selection was having none of it. Anyway, Vintana overcame this craniofacial adversity with characteristic Mesozoic moxie, and is an important piece in the puzzle of mammal evolution. It will be interesting to see what other mammalian surprises the Mesozoic has in store for paleontologists.

eFfing Fossil Friday: Frozen Femur

A 45,000 year old human femur from Siberia provides new information about genetic mutation rates and modern human origins. As Quiaomei Fu and colleagues report in this week’s issue of Nature, this seemingly simple leg bone carries so much information, not because of its gross anatomy, but because of the ancient DNA it preserves.

The femur wasn’t discovered by paleontologists, but by an artist/historian looking for fossils around the Irtysh River. The bone came from from a site called Ust’-Ishim, only some 650 km north of the snowy capital where I work in Kazakhstan:


The site in question, Ust’-Ishim is marked by the yellow star. The red and blue sites to the southeast are other Upper Paleolithic sites. Okladnikov (3) and Denisova (4) have also yielded fossils preserving ancient DNA. Modified from Fu et al. figure 1.

The bone was directly radiocarbon dated to around 45,000 years ago. With a fairly precise age of the bone, Fu et al. could estimate the rate at which genetic mutations arise, by counting the number of new mutations in recent humans that aren’t shared by the Ust’-Ishim femur. This led to an estimate of around 0.43×10−9  new mutations per site per year. This is a relatively low rate compared to estimates based on geologically older fossils, but consistent with more recent estimates that directly compare parents and offspring.

The Ust’-Ishim individual had levels of Neandertal ancestry comparable to living Eurasians (~2.3% of the genome), but there is no evidence of any Denisovan ancestry. Because this individual lived closer to the date of modern-Neandertal admixture, the Neandertal segments of its genome are longer than in modern people (recombination over generations breaks these regions apart into shorter segments). Knowing about recombination rates, Fu et al. could infer that admixture between Neandertal and modern human populations occurred between 50-60,000 years ago.

This eFfing Friday fossil provides more tantalizing evidence for DNA-bearing human fossils just across the Kazakhstan border. With Ust’-Ishim to the north, Denisova and Okladnikov caves to the east, and Teshik Tash to the south, my colleagues and I are very keen to find similar sites here on the KZ side.

Reference: Fu et al. 2014. Genome sequence of a 45,000-year-old modern human from Siberia. Nature 514: 445–449. doi:10.1038/nature13810.