Another small Middle Pleistocene person

Last year I brought up the implications of the small female pelvis from Gona, Ethiopia for body size variation in Homo erectus (see previous post). This individual was much smaller than other Middle Pleistocene Homo fossils, indicating size variation comparable to highly sexually dimorphic gorillas and unlike recent human populations. Before this pelvis, most known Homo erectus fossils were fairly large (comparable to living people), with only a few hints of much smaller individuals (e.g., KNM-ER 427000, KNM-OL 45500). Now joining this petite party, this tiny troop, this little lot, this compact cadre, etc., is KNM-WT 51261, a 750,000 year old molar from Kenya (Maddux et al., in press).

Occlusal area for hominin first molars. The tooth is from Fig. 2 and the plot from Fig. 3 in the paper.

Occlusal area for first molars in the genus Homo. The tooth image is from Fig. 2 and the plot from Fig. 3 in Maddux et al. Lookit how tiny it is!

This ‘new’ specimen substantially increases the range of size variation among early African H. erectus molars, although the expanded range isn’t remarkable compared with later Homo samples such as from Zhoukoudian cave in China or Neandertals. What is different, though, is that most of the highly variable samples show a fairly continuous range of variation, while the WT 51261 molar is a considerable outlier from the rest of the African Middle Pleistocene sample (a lot like the situation with the Gona pelvis). So this tooth re-raises an important question: were smaller specimens like Gona and WT 51261 as rare in life as they are in the fossil record, or was such great size variation common in the Middle Pleistocene? How we reconstruct what kind of animal Homo erectus was differs depending on the answer to this question.

Driving nails into the 2014 Lawn Chair

It’s that time again, when we come to bury the year we’ve just defeated, only to celebrate the zombie birth of a new onslaught of days to clobber. In the spirit of auld lang syne, let’s recap the highlights of Lawn Chair in 2014.Georgia dinos 2014

Osteology was everywhere: although I am wont to see bones everywhere in everyday life, this year I only wrote about it four times. First there were the baby bones in cafe upholstery in my hometown of Kansas City, then the giant sheep bones in my new home of Astana. I discovered that animal bones littered the landscape of desert Mangystau, and then I spotted a vertebra hiding in a helmet at a conference in Italy. I also tweeted about a false femur head from a karaoke bar in Astana. You can’t escape. 2015 is sure to be more osseous.BONES!
eFfing Fossil Friday reboot: This old series focusing on fossils furtively restarted on a plane, when I uncovered the conspiracy that the Australopithecus africanus cranium Sts 71 was actually the Kryptonian codex. I later wrote about the Sima de los Huesos skulls, Neandertal poop, the origins of feathers on badass dinosaurs, the 45,000 year old Ust’-Ishim femur and its delicious DNA, and facial flanges in early mammals and nearly modern baboons. Fossils are the best, and 2015 is bound to be as fossiliferous as last year.Ancient DNA was boss: In addition to the earliest ‘modern’ human DNA from Ust’-Ishim, 2014 also witnessed a swath of studies early on attesting to the success of paleogenomics. We also got a first glimpse into epigenetics of ancient humans, and the potential importance this will have in uncovering how our DNA makes us human. Along these lines, for 2015, I’d be keen to see more work on miRNA and other aspects of gene regulation in ancient genomes.

Screen Shot 2014-10-24 at 11.26.31 AMR codes: I’ve posted R code for the analysis from my paper that came out this year, comparing mandibular growth in humans and Australopithecus robustus (I didn’t get to talk about that paper when it came out because I was in the middle of the Rising Star Workshop. Things to look forward to in 2015…). I’ll also be posting code for the analysis of brain growth in Homo erectus once that paper is published, and I have already posted code for creating the pretty pictures from the paper.

Brain size data (left) and the average annual rates from birth calculated from pairs of specimens (right). Black=humans, green=chimpanzees, red=gorillas, blue=Homo erectus.
Body size variation in Homo erectus: A response to a response to a paper led me to reexamine sexual dimorphism in body size in our early ancestor – seems it was higher than has lately been appreciated, and there are many potential reasons for this. I presented the initial results of this investigation on the blog and at a conference, and am now writing this up for publication. This investigation is based on resampling statistics, nothing as new and flashy as in the growth studies. I will post code for these analyses on the R Codes page in due time.

Dimorphism ratios copy

Resampled ratios of dimorphism, calculated by dividing the average of six randomly selected male body masses by a randomly selected female mass. The blue star in each plot is the empirical ratio of average male mass/average female mass. For all species the average resampled ratio is almost identical to this empirical value. The red line marks the ratio of the six largest (male?) Homo erectus mass estimates divided by the estimated mass of the Gona (female?) pelvis. The Homo erectus male/female difference is rarely observed in chimps and humans, but is common in gorillas. Gorillas display high levels of sexual dimorphism, suggesting this may have been the case for Homo erectus as well.

Classroom lab activities: This year I added a lab components to my courses here at NU, and I posted up two of the lab activities I did in my classes this semester. Last spring, I got the idea for an activity in which students measure toe joint angles on digital images, to test whether Ardipithecus kadabba and other hominin toes can be distinguished from apes’. This semester, students in my human evo-devo class did this study, and generally found hominin toes to be more angled than apes’. Hypothesis tested. My Intro to Bio Anthro class tested whether their limb proportions fit expectations based on Allen’s Rule, and mystery ensued. My classes next term aren’t as conducive to lab activities, but if I come up with any good assignments I’ll be sure to post them.class models both copy

Now that 2014 is laid to rest, here’s to a bright and successful zombied 2015! Жаңа Жылыңызбен!

2015 AAPA conference: More brain growth

The American Association of Physical Anthropologists is holding its annual meeting next year in St. Louis, in my home state of Missouri (I’m from Kansas City, which is by far the best city in the state, if not the entirety of the Midwest). I’ll be giving a talk comparing brain size growth in captive and wild chimpanzees, on Saturday 28 March in the Primate Life History session. Here’s a sneak peak:

Velocity curve for brain size from birth to 5 years in wild (green) and caprive (blue) chimpanzees. For the captive models, the dashed line is fit to the raw brain masses, and the solid line is fit to the estimated endocranial volumes.

Velocity curves for brain size growth from birth to 5 years in wild (green) and captive (blue) chimpanzees. The wild data are endocranial volumes, but the captive specimens are represented by brain masses. So the captive data are modeled for both the original masses (dashed) and estimated volumes (solid). Wild data are from Neubauer et al. 2011, captive data from Herndon et al., 1999.

Abstract: This study compares postnatal brain size change in two important chimpanzee samples: brain masses of captive apes at the Yerkes National Primate Research Center, and endocranial volumes (ECVs) of wild-collected individuals from the Taï Forest. Importantly, age at death is known for every individual, so these cross-sectional samples allow inferences of patterns and rates of brain growth in these populations. Previous studies have revealed differences in growth and health between wild and captive animals, but such habitat effects have yet to be investigated for brain growth. It has also been hypothesized that brain mass and endocranial volume follow different growth curves. To address these issues, I compare the Yerkes brain mass data (n=70) with the Taï ECVs (n=30), modeling both size and velocity change over time with polynomial regression. Yerkes masses overlap with Taï volumes at all ages, though values for the former tend to be slightly elevated over the latter. Velocity curves indicate that growth decelerates more rapidly for mass than ECV. Both velocity curves come to encompass zero between three and four years of age, with Yerkes mass slightly preceding Taï ECV. Thus, Yerkes brain masses and Taï ECVs show a very similar pattern of size change, but there are minor differences indicating at least a small effect of differences in habitat, unit of measurement, or a combination of both. The overall similarity between datasets, however, points to the canalization of brain growth in Pan troglodytes.

Kazakhstan’s killer cats

I’m reading up on previous paleontological research that’s taken place here in Kazakhstan, planning for future work. There aren’t any human fossils known from here (at least, none to my knowledge). But, I did stumble upon this badass, sabre-toothed cat from the Late Miocene (over 5 million years ago):

From Sotnikova, 1992. Original caption: Fig. 2. Machairodus kurteni, Kalmakpai (PIN-2433/287), skull, ventral and lateral view.

From Sotnikova, 1992 (mandible not shown). Original caption: Fig. 2. Machairodus kurteni, Kalmakpai (PIN-2433/287), skull, ventral and lateral view.

The skull was described by MV Sotnikova in 1992, and comes from a site called Kalmakpai in the Zaysan Basin in East Kazakhstan. For perspective, Sotnikova says the skull is about the same size as an adult African lion. This is much larger than the wildlife I’ve seen lately in snow-soaked Astana (I trailed a large white rabbit in Presidential Park by the river on my run today. Not as badass).

This is a reminder that the Big Cats once had a much larger geographical distribution than they do today. The skull above belongs to the genus Machairodus, which is also known from Africa, Europe and North America. Machairodus is closely related to Homotherium, another large, geographically dispersed genus of sabre-toothed cat from the Pliocene (including at Dmanisi).

Of course, extinction isn’t exclusive to the deep past: the Caspian Tiger used to roam parts of southern Kazakhstan and other areas of Central Asia, going extinct only in the past few decades.

Results of the toe-tally easy lab activity

Alternate title: Dorsal canting in primate PPP4s

Earlier this year I suggested a classroom activity in which students can scrutinize the evidence used to argue that the >5 million year old (mya) Ardipithecus kadabba was bipedal. To recap: Ar. kadabba is represented by some teeth, a broken lower jaw, and some fragmentary postcrania. The main piece of evidence that it is a human ancestor and not just any old ape is from a single toe bone, and the orientation of its proximal joint. In Ar. kadabba and animals that hyperdorxiflex their toes (i.e., humans and other bipeds when walking), this joint faces upward, whereas it points backward or even downward in apes. This “dorsal canting” of the proximal toe joint has also been used as evidence that the 4.4 mya Ardipithecus ramidus and 3.5 mya owner of the mystery foot from Burtele are bipedal hominins. A question remains, though – does this anatomy really distinguish locomotor groups such as bipeds from quadrupeds?

Use ImageJ to measure the canting angle between the proximal joint and plantar surface. Proximal to the right, distal to the left.

STUDENT SCIENTISTS TO THE RESCUE! Use ImageJ to measure the canting angle between the proximal joint and plantar surface, as I’ve done on this Japanese macaque monkey (they are not bipedal). Proximal to the right, distal to the left Note I changed the measured angle from the March post.

I sicked my students in Ant 364 (Human Evolutionary Developmental Biology) here at NU on this task. I had students look at only 11 modern primates from the awesome KUPRI database. Most groups are only represented by 1 (Homo sapiens, Hylobates lar and Macaca fuscata) or two (Pongo species and Gorilla gorilla) specimens, all adults. For chimpanzees (Pan troglodytes) there is one infant and four adults. The database has more individuals, and it would be better to include more specimens to get better ideas of species’ ranges of variation, but this is a good training sample for a class assignment. The fossil group includes one Ardipithecus ramidus, one Ar. kadabba, one Australopithecus afarensis, and the PPP4 of the mystery foot from Burtele. The human and all fossils except Ar. kadabba are based off of lateral photographs and not CT scans like for the living primates, meaning there may be some error in their measurements, but we’ll assume for the assignment this is not a problem. Here are their results:

Dorsal canting angle of the fourth proximal pedal phalanx in primates.

Dorsal canting angle of the fourth proximal pedal phalanx in primates. The lower the angle, the more dorsally canted the proximal joint surface. The “Fossil” group includes specimens attributed to ArdipithecusAustralopithecus and something unknown.

Great apes have fairly high angles, meaning generally not dorsally canted proximal joint surfaces. The two gorillas fall right in the adult chimpanzee (adult) range of variation, while chimp infant and orangutans have much higher angles (≥90º means they’re actually angled downward or plantarly). The gibbon (Hylobates) is slightly lower than the chimpanzee range. The macaque has an even more dorsally canted joint, and the human even more so. The fossils, except the measurement for Ar. ramidus (see note above), have lower angles than living apes, but higher than the human and the monkey. If dorsal canting really is really a bony adaptation to forces experienced during life, then the fossil angles suggest these animals’ toes were dorsiflexed more so than living great apes (but not as much as the single monkey and human).

This lab helps students become familiar with CT data, the fossil record, taking measurements (students also measure maximum length of the toe bones and look at the relationship between length and canting), analyzing data, and hypothesis testing. You can also have fun exploring inter-observer error by comparing students’ measurements.

Here’s the full lab handout if you want to use or modify it for your own class: Lab 5-Toe instructions and report

Lessons from limb lab (activity)

This semester I have added a lab component to my Introduction to Biological Anthropology class. Lab activities and assignments provide students with opportunities to gather data, to think about them in the context of various theories, and to learn about how to analyze them. This past week’s lab looked at limb proportions within our classroom, in the context of “Allen’s rule” – in colder climates, animals tend to have relatively shorter distal limbs (radius+ulna and tibia+fibula). Allen’s rule, along with “Bergmann’s rule,” describe ecogeographic variation in humans and other animals: body size (i.e., mass) and shape (i.e., limb proportions) tend to vary with climate, such that populations living in colder environments tend to have less surface area relative to body mass, as an adapation to retain body heat.

A simple and effective way to quantify the relative lengths of limbs is through ratios: in this case we examined the crural and brachial indices. Here I’ve plotted average human indices against latitude as reported in a recent paper by Helen Kurki and colleagues (2008):

Left: Crural index (tibia length/femur length) related to latitude, as a proxy for climate. Right: Brachial index (radius length/humerus length) plotted against latitude. Data from Kurki et al. (2008). Black=female, red=male. Dashed lines are  regression slopes for each sex, and solid lines indicate the 95% confidence limits of the regression lines.

Human limb proportions related to latitude, as a proxy for climate. Left: Crural index (tibia length/femur length). Right: Brachial index (radius length/humerus length). Higher indices mean relatively longer tibia or radius. Data from Kurki et al. (2008). Black=female, red=male. Dashed lines are least squares regression lines for each sex, and solid lines indicate the 95% confidence limits of the regression lines. How well will our class’s data fit these models?…

These plots are consistent with Allen’s rule – the distal limb segments become relatively shorter with increasing latitude. In lab, we test whether our limb proportions reflect this presumably ecogeographic pattern. Here in Astana we are at 51ºN latitude, so these regressions predict that our class should have crural indices between 0.82-0.84, and brachial indices between 0.72-0.79. How well does our class fit these model’s predictions?

Pretty terribly.

…Pretty terribly. Plots are same as above, except with our class’s data added at 51º N latitude (vertical lines). In each, the vertical lines span the class’s 95% range (black=females, red=males), with the dots marking each sex’s average. Kazakhstan is huge, and students could have grown up in latitudes from 42º-55º N, but even assuming students had all come from Shymkent their distal limbs still appear much longer than expected.

These plots show that students in the class have longer distal limbs than expected – both for our latitude, and for humans generally. The poor fit of my students’ limb proportions probably doesn’t mean they’re bad humans. Instead, we probably deviate because we compared apples to oranges: the Kurki data were dry long bones measured on an osteometric board, whereas I had my students do their best to palpate and measure the maximum lengths of their own bones beneath layers of fat, muscle, skin, and clothing. Our high indices probably reflect the underestimation of humerus and femur lengths, whose most proximal points that can be palapated (greater tubercle and trochanter, respectively) lie a bit lower than the respective heads, which would have been included in the Kurki measurements.

It was interesting to review these plots with students. Even though they’re fairly new at reading graphs like these, there was an audible gasp and bewildered muttering when their own data went up on the board. I myself was surprised at these results, but I’m happy with how the exercise went. This particular ‘study’ helps students learn about ecogeography, adaptation and human variation, as well as the importance of homology and comparing like with like.

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.

Can ‘ape-like’ actually be ‘human-like’?

I’m reading up on life history in Homo erectus for a few projects I’m working on, and something’s just caught my eye. A 2012 issue of Current Anthropology presents a series of papers from the 2011 symposium, “Human Biology and the Origins of Homo.” This issue is full of great stuff, and to top it all off, it can be accessed online for free! (here’s the JSTOR link)

Gary Schwartz has a paper here recounting what is known (or as he stresses, what is still largely unknown) about growth and life history in early Homo. Dental evidence accumulated over the past 30 years has pointed to a rapid (ape-like) life cycle for fossil hominins, in comparison with a slow, long and drawn out human pattern. But much of the evidence against a human-like pattern is somewhat indirect. For instance, Holly Smith (1991) has shown that there’s a pretty tight relationship between brain size and age at first molar (M1) eruption in Primates:

M1 crancap

Fig. 1 from Schwartz (2012). “Bivariate plot of ln M1 emergence age in months (y) versus ln cranial capacity in cubic centimeters (x) for a sample of anthropoids.” The hominins and humans are the open shapes, to which I’ve visually fitted the red line.

It’s a very high correlation (r=0.98). This means that armed with simply an animal’s brain size (“cranial capacity” in the graph), which is fairly easy to estimate given complete enough fossils, one can estimate with a bit of confidence its likely age range for M1 emergence. With brain sizes between apes’ and ours, fossil hominins can be estimated to have erupted their M1s at younger ages than us. Many subsequent studies of tooth formation, based on the microscopic remnants of tooth development, have supported these inferences. So presumably, faster, ape-like dental development could be extrapolated to mean ape-like body growth rates and other aspects of life history as well.

But although this is a tight relationship, there are deviations. As Schwartz notes in the article, and others have noted before, high correlations found when examining large interspecific groups (e.g., primates as a whole) often break down when the focus is on smaller groups of more closely related species (e.g., just apes). Based on the relationship figured above, humans are expected to erupt M1 around 7 years of age, but nearly all humans erupt M1 closer to 6 years (hence the open diamond for humans is below the regression line). What hominins appear to share in common with humans is a younger age at M1 eruption than expected for primates of their brain sizes (the red line I’ve added to the figure).

Hominins’ faster dental development and eruption may be ape-like in absolute terms, but eruption ages may be human-like when their brain size is taken to account. As with many life history variables, the significance of this similarity (if anything) is difficult to ascertain.

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:

Ust'-Ishim

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.

Osteology everywhere: Pollicem verte(b)r(a)e [Latin puns are hard]

I just got back from the meetings of the European Society for the Study of Human Evolution in Florence. As you can guess, bones and genes and anatomy and apes and biomechanics and energetics and everything were on everyone’s minds. Even in the midst of an unseasonal surprise typhoon of lunch time ice:

Ambush of hail.

Aw hail no.

Along the way, I passed a gift shop window and this book cover immediately caught my eye:helert

No, it’s not an ancient Roman gladiator’s helmet. It’s clearly a lumbar vertebra, probably of some quadruped. We’re looking down onto the top (or front of it) from the cranial view. The body or centrum is the rounded part toward the bottom of the picture, the short transverse processes jutting off to the sides. The spinous process, pointing toward the top, is even thick and blunt distally as is characteristic of lumbar verts. Here’s a comparison:

Middle lumbar vertebrae, from the cranial view (modified from Figs. 3-4 of Moyà-Solà et al., 2004). 0=modern baboon, A=Proconsul nyanzae (KNM-MW 13142-J)(B) P. catalaunicus (IPS-21350.59). (C) Cast of Morotopithecus bishopi (UPM 67.28) from Moroto (Uganda). (D) D. laietanus (IPS-18000) from Can Llobateres (Spain). (E) Pongo pygmaeus

Middle lumbar vertebrae of various Miocene apes (A-D) in cranial view (modified from Figs. 3-4 of Moyà-Solà et al., 2004). 0=modern baboon, A=Proconsul nyanzae (KNM-MW 13142-J), B=Pierolapithecus catalaunicus (IPS-21350.59), C=Morotopithecus bishopi (UPM 67.28), D=Hispanopithecus laietanus (IPS-18000), and E= modern orangutan.

Modern apes use an upright posture more frequently than living monkeys, who are quadrupedal. An anatomical correlate of these postures is the position of the transverse processes. Compare the baboon (0 in the figure above) with the orangutan (E). In the monkey the transverse processes come off the sides of the centrum (below the horizontal line), while in the orangutan the processes come off the pedicle further back. In your lumbars the transverse processes arise a little bit more toward the back than in the orangutan.

This is a pretty characteristic pattern, meaning that we can reconstruct the habitual posture of an animal based on a single bone – even just part of a single bone as in the case of Hispanopithecus (D, above). Proconsul nyanzae (A), dating to around 19 million years ago and therefore one of the earliest apes, has a monkey-like lumbar vert; the rest of its skeleton is monkey-like and so we think many of the earliest apes moved around like modern monkeys. In contrast, Morotopithecus bishopi (C), at 20.6 million years ago, is also one of the earliest apes but has a more modern-ape-like lumbar. And so with Pierolapithecus and Hispanopithecus.

The vertebra gracing the cover of our gift shop book is clearly more monkey-like, presumably from a simian who long ago walked on all fours across the blood-soaked floors of a cacophonous Colosseum.