How small apes grow big canine teeth

Gibbons are sometimes referred to as “lesser apes” since they’re the smaller-bodied cousins of “great apes” like us humans, chimpanzees, gorillas, and orangutans. But what they lack in body mass they make up for in taxonomic diversity, with roughly 20 species distributed across four genus groups (Kim et al., 2011). And while male great apes (except humans) have large canine teeth, both sexes in gibbons have large maxillary canines — flashy weaponry for defending territory.

Pointy canine teeth peeking out from the upper and lower jaws of an adult female gibbon cared for at the International Primate Protection League (source)

My research has generally focused on brains and growth throughout human evolution, but I started looking at gibbons a few years ago when the COVID-19 pandemic put research travel on hold. Inspired by Julia Zichello’s 2018 article about gibbon models for understanding hominin evolution and appreciating that “overlooked small apes need more attention,” I had the opportunity to CT scan a unique skeletal collection of white-handed gibbons (Hylobates lar), which was sadly harvested from the forests of Thailand back in the late 1930s. Previous research on skull growth in gibbons has mostly used small samples compiled from different species (and sometimes even different genera). In contrast, this CT dataset includes many individuals at each stage of maturation from late infancy through adulthood, effectively representing a single population at a point in time. So with this larger cross-sectional sample of a single species, we can better understand how gibbon brains and faces grow. And because permanent teeth form in a long, continuous sequence throughout the growth period, an individual’s state of dental development can serve as a marker of where they are along the maturation process.

In a paper hot off the press, Julia Boughner and I analyzed dental development in this unique sample (article here). One of the coolest things we found was that gibbons’ large upper canine teeth are among the first to begin but last to finish tooth formation. In fact, the large canines growing inside relatively small faces may inhibit growth of one of the neighboring incisor teeth until the face has grown to create enough space for it. And while most teeth developing within the jaw begin emerging into the mouth once there’s enough room for them, gibbons’ gargantuan upper canines are forced out of hiding as they outgrow their bony crypts (check out the right-most jaw in the second row below).

Cross-sectional representation of tooth formation in white-handed gibbons, starting with the youngest in the top left and ending with the oldest in the bottom right. The first permanent tooth to form and emerge, M1, is highlighted along with the canine “C.”

In addition to characterizing ‘normal’ dental development, we also observed several developmental anomalies and pathologies in the sample. Our observations corroborate previous research showing that tooth formation generally proceeds ‘as scheduled’ despite various other disturbances to development.

It remains to be seen whether early development of the canine at the cost of delayed incisor formation is a pattern unique among all the apes, since most other studies of ape tooth formation have examined the lower jaw while our study focused on the upper jaws. But the canine-incisor tradeoff that we identified sets the stage for subsequent study of skull growth in this sample, as it highlights the many factors and functions that must be coordinated during growth.

While we have several projects planned with this unique dataset, we have also published the tooth formation data that we analyzed, and the original micro-CT scans themselves will be published to the online repository Morphosource.org soon, once a few more projects are finished.

Worst year in review

As we’re wrapping up what may be the worst year in recent global memory, especially geopolitically, let’s take a moment to review some more positive things that came up at Lawnchair in 2016.

Headed home

IMG_20160801_143520

Alternate subtitle: Go West
This was a quiet year on the blog, with only 18 posts compared with the roughly thirty per year in 2014-2015. The major reason for the silence was that I moved from Kazakhstan back to the US to join the Anthropology Department at Vassar College in New York. With all the movement there was  less time to blog. Much of the second half of 2016 was spent setting up the Biological Anthropology Lab at Vassar, which will focus on “virtual” anthropology, including 3D surface scanning…

20161111_141908.jpg

Cast of early Homo cranium KNM-ER 1470 and 3D surface scan made in the lab using an Artec Spider.

… and 3D printing.

20161107_135616

gibbon endocast, created from a CT scan using Avizo software and printed on a Zortrax M200.

This first semester stateside I reworked my ‘Intro to Bio Anthro’ and ‘Race’ courses, which I think went pretty well being presented to an American audience for the first time. The latter class examines human biological variation, situating empirical observations in modern and historical social contexts. This is an especially important class today as 2016 saw a rise in nationalist and racist movements across the globe. Just yesterday Sarah Zhang published an essay in The Atlantic titled, “Will the Alt-right peddle a new kind of racist genetics?” It’s a great read, and I’m pleased to say that in the Race class this semester, we addressed all of the various social and scientific issues that came up in that piece. Admittedly though, I’m dismayed that this scary question has to be raised at this point in time, but it’s important for scholars to address and publicize given our society’s tragically short and selective memory.

So the first semester went well, and next semester I’ll be teaching a seminar focused on Homo naledi and a mid-level course on the prehistory of Central Asia. The Homo naledi class will be lots of fun, as we’ll used 3D printouts of H. naledi and other hominin species to address questions in human evolution. The Central Asia class will be good prep for when I return to Kazakhstan next summer to continue the hunt for human fossils in the country.

Osteology is still everywhere

A recurring segment over the years has been “Osteology Everywhere,” in which I recount how something I’ve seen out and about reminds me of a certain bone or fossil. Five of the blog 18 posts this year were OAs, and four of these were fossiliferous: I saw …

2016-02-09 16.26.31

Anatomy terminology hidden in 3D block letters,

Picture1

Hominin canines in Kazakhstani baursaki cakes,

Picture1

The Ardipithecus ramidus ilium in Almaty,

screen-shot-2016-11-26-at-5-36-05-pm

Homo naledi juvenile femur head in nutmeg,

bangkok-erectus

And a Homo erectus cranium on a Bangkok sidewalk. As I’m teaching a fossil-focused seminar next semester, OA will probably become increasingly about fossils, and I’ll probably get my students involved in the fun as well.

New discoveries and enduring questions

The most-read post on the blog this year was about the recovery of the oldest human Nuclear DNA, from the 450,000 year old Sima de los Huesos fossils. My 2013 prediction that nuclear DNA would conflict with mtDNA by showing these hominins to be closer to Neandertals than Denisovans was shown to be correct.

giphy

These results are significant in part because they demonstrate one way that new insights can be gained from fossils that have been known for years. But more intriguingly, the ability of researchers to extract DNA from exceedingly old fossils suggests that this is only the tip of the iceberg.

The other major discoveries I covered this year were the capuchin monkeys who made stone tools and the possibility that living humans and extinct Neandertals share a common pattern of brain development.

Pride & Predator

An unrelated image from 2016 that makes me laugh.

The comparison between monkey-made and anthropogenic stone tools drives home the now dated fact that humans aren’t the only rock-modifiers. But the significance for the evolution of human tool use is less clear cut – what are the parallels (if any) in the motivation and modification of rocks between hominins and capuchins, who haven’t shared a common ancestor for tens of millions of years? I’m sure we’ll hear more on that in the coming years.

In the case of whether Neandertal brain development is like that of humans, I pointed out that new study’s results differ from previous research probably because of differences samples and methods. The only way to reconcile this issue is for the two teams of researchers, one based in Zurich and the other in Leipzig, to come together or for a third party to try their hand at the analysis. Maybe we’ll see this in 2017, maybe not.

There were other cool things in 2016 that I just didn’t get around to writing about, such as the publication of new Laetoli footprints with accompanying free 3D scans, new papers on Homo naledi that are in press in the Journal of Human Evolution, and new analysis of old Lucy (Australopithecus afarensis) fossils suggesting that she spent a lifetime climbing trees but may have sucked at it. But here’s hoping that 2017 tops 2016, on the blog, in the fossil record, and basically on Earth in general.

Arm and leg modelling

No, I’m not looking for people with lithe limbs to be photographed for money. Much more sexily, I’m referring to a recent paper (Pietak et al., 2013) that’s found that the relative length of the segments of human limbs can be modeled with a log-periodic function:

Figure 2 from Pietak et al. 2013. Human within-limb proportions are such that the length of each segment (e.g., H1-6) of a limb, from  fingertip to shoulder (A) and to to hip (B), can be predicted by a logarithmic periodic function (C).

In other words, within a limb, the length of each segment is mathematically fairly predictable on the basis of the segment(s) before and after it. As the authors state, “Being able to describe human limb bone lengths in terms of a log-periodic function means that only one parameter, the wavelength λ, is needed to explain the proportional configuration of the limb.”

The biological significance of this pattern is difficult to discern. The length of a limb segment is determined by a number of factors, including the spacing between the initial limb condensations embryonically, and thereafter the growth rates and duration of growth at proximal and distal epiphyses. As a result, limb proportions aren’t static throughout life, but change from embryo to adult. For instance, here are limb proportion data for the coolest animal ever – gibbons! – from the great anatomist Adolph Schultz.

ResearchBlogging.orgAn important question, and follow-up to Pietak et al’s study, is whether human limb proportions can be described by such log-periodic functions throughout ontogeny, and if so how these change. Plus, it’s also not clear to what extent human proportions might happen to be describable by log periodic functions, simply because each segment is shorter than the one preceding it proximally. In short, this study raises really interesting and pursuable questions about how and why animal limbs grow to the size and proportions that they do.

References
Pietak A, Ma S, Beck CW, & Stringer MD (2013). Fundamental ratios and logarithmic periodicity in human limb bones. Journal of anatomy, 222 (5), 526-37 PMID: 23521756

Schultz, A. (1944). Age changes and variability in gibbons. A Morphological study on a population sample of a man-like ape American Journal of Physical Anthropology, 2 (1), 1-129 DOI: 10.1002/ajpa.1330020102