Is Homo floresiensis really that strange?

BMC Biology has recently published a paper (It’s Open Access!) which explores trends in brain size in the Primates.  A trend toward a larger brain is usually considered one of the “hallmarks” of the Primates, but Stephen Montgomery and his colleagues have shown that in many lineages, there is a trend towards secondarily “shrunken” brains.

The authors looked at three different traits- absolute brain mass, absolute body mass, and relative brain mass (a derivitive of brain mass and body mass) in 37 living species, and 23 extinct species.  They reconstructed the ancestral state using three different phylogenetic methods:  Parsimony, maximum likelihood, and a Bayesian Markov-chain Monte Carlo.  They found that there are increases in both absolute and relative brain size in the Primate lineage, but not necessarily in body size.  So primates, in general, have more brain per pound of body than most other mammals.

In a few branches, once an initial increase in brain size occurred, there was a secondary decrease in brain size.  On the surface, it would seem that this doesn’t make sense.  More brain= more smarts= more behavioral flexibility= more food, more mating, more survival.  Right?  Usually.  But in some species, the energy it costs to maintain all of that extra brain costs more than it’s worth.  If you are a bat who hunts for insects, you may not need to remember the location of all of the fruiting trees in the area, and the extra weight incurred by having a big brain weighs you down when you’re trying to fly.  In that case, it might be advantageous to reduce your brain size.

There’s also the issue of a “phyletic dwarf” or “phyletic giant.”  These are species which are very closely related to each other, but one is very big or very small.  Something like the Aye-aye and the Giant Aye-aye.  Brain size generally exhibits negative allometry because it’s such a specialized organ- so, as the body gets larger, the brain doesn’t keep up and is, as a result, proportionately smaller.

It’s extremely important for most of your organs to increase with body size.  For example, a bigger animal needs to pump more blood, so it needs a bigger heart.  A bigger animal eats more food and needs a bigger liver.  There are certain areas of the brain that increase allometrically with body size- usually areas that are in charge of motor skills.  If you’ve got bigger legs, you’ve got bigger muscles, and you need more neural projections in order to control them.  But does a larger animal need to think more?  Will it benefit from an extra few cubic centimeters of neocortex?  Probably not, so it’s not worth the extra time and energy it takes to develop that neocortex.

And that sort of brings us to an important question in evolutionary neurobiology: Does absolute brain size matter, or is it solely brain size relative to body size?  Brains that are absolutely larger have more neurons, which could have important cognitive implications.  But how many of those extra neurons are just being used to control the physiological functions of the body?

Does size even tell us anything at all?  Any way you look at it, brain size is a crude measurement of cognitive ability.  In an ideal world, we would know the proportion of each of the different regions of the brain in each species and go from there.  But, those kinds of measurements are hard to obtain in living species, and impossible in fossils.  Ralph Holloway has been saying since 1967 that there has got to be a better way than just plain ol’ cranial capacity… but other than noting the relative position of different sulci and gyri on endocasts, there isn’t too much else to be done.

Anyway, those considerations aside, Montgomery and friends found that there were initial increases in proportional brain size- one at the node between ancestral primates and strepsirrhines, another between ancestral primates and haplorrhines, and then another between the ancestral haplorrhines and anthropoids.  At the terminal ends of the branch, tarsiers, galagos, aye-ayes, and humans all show large increases in relative brain size.

Cool!  But what about the decreases in brain size?  They found that absolute brain size decreased in about 14% of the branches, in clades like Mangabeys, tamarins and marmosets, and some of the small lemurs.  In every one of those cases, the decrease in brain mass was accompanied by a decrease in body mass.  And body mass decreased a lot more often than brain mass- 46% of the branches showed a decrease in body mass.  So what that means is that brains tend to stay the same size in lineages where the body size is decreasing.

Only 4% of the branches showed a decrease in relative brain size.  Most of these are lineages where the body size increased disproportionately to brain size- the negative allometry discussed above.  Think about gorillas:  They have a small proportion of brain mass to body mass.  But is it because they have tiny brains, or because they have huge bodies?  This study seems to support the idea that it’s their large bodies that are influencing the numbers.

Okay, so we’ve got lots of increases in brain size, and a few decreases.  In the cases where we have decreases, we usually have body size decreases as well.  More often than not, we have body size decreases which result in a disproportionately large brain size, but occasionally we have a body size increase which results in a disproportionately small brain size.  And all of that brings us to the Hobbit.

The authors looked at Homo floresiensis in relation to the Dmanisi hominids, Homo habilis, and a Homo erectus from Ngangdong and found that if we use Dmanisi or habilis as an ancestor, the decrease in brain size and body size isn’t exceptionally weird when compared to other primate groups.  The mouse lemur decreased in both to a greater degree, for example.

But if you use the Ngangdong erectus as the ancestor, it is a really weird decrease.

So, I guess the question is, is it reasonable to use Dmanisi or Homo habilis as the ancestor and not Homo erectus?  And of course, we don’t know that yet!

Montgomery, S., Capellini, I., Barton, R., & Mundy, N. (2010). Reconstructing the ups and downs of primate brain evolution: implications for adaptive hypotheses and Homo floresiensis BMC Biology, 8 (1) DOI: 10.1186/1741-7007-8-9

Sapolsky on TED talks

Robert Sapolsky. ‘Nuff said.

Okay, maybe I will say one more thing: Sapolsky’s book A Primate’s Memoir is one of the most beautiful books I have ever read, and I really don’t think it’s because I am a primate geek. I think it’s one of the rare science books that would be enjoyable for anyone, or maybe everyone, to read.

Four Stone Hearth #84! (Gratuitous Gelada Edition)

Welcome to the 84th edition of the Four Stone Hearth!  The last time I did this, I tried to separate the posts into whichever “subfield” they fit into, but the intersectionality this time was phenomenal.  Grab your favorite beverage and sit down for some good old-fashioned anthropology reading.

It’s not very often that we get to talk about culture from primates other than ourselves (though it’s becoming more and more frequent!), but Eric Michael Johnson discusses Susan Savage-Rumbaugh’s TED talk on Bonobos and the emergence of culture at The Primate Diaries.

Speaking of chimps, Michelle at SpiderMonkeyTales writes about the Fongoli Chimps and the things they do to cope with a more marginal environment.  Nocturnal Chimps?  In addition, the same chimps can understand fire.

Continuing on with the chimps-in-marginal-environments theme, Greg Laden discusses chimps, underground storage organs, volcanoes, and what all of them have to do with human evolution over at his place.

Raymond Ho writes about obedient young Campbell’s monkeys over at The Prancing Papio.

Beast Ape has written about Friendship, Fatherhood, and MHC in baboons.  They’ve done lots of stuff with mate choice and MHC, but this is the first time I’ve seen it used in a parent-offspring project- with surprising results!  Really, really cool.

Speaking of the bleeding heart baboons, I revealed that I have a major crush on geladas here.  And why am I so sure that knuckle-walking evolved twice?  And is it tacky to link to your own posts when you’re hosting a blog carnival?

Anna has written a beautiful piece about Childhood- its evolutionary origins, its philosophical implications, and cross-cultural variability- in The Ape That Wouldn’t Grow Up.  Happy birthday, Anna!

Zacharoo from Lawn Chair Anthropology has written a fascinating post about signatures of hybridization in gorillas.  Neat stuff!

On the more human side of things, Tim Jones at anthropology.net draws our attention to a really cool skeletal pathology found at Atapueurca- a craniosyntosis!- and what it means (or what it might not mean) about the sociobiology of Pleistocene hominids.

And, while he was at it, he reviewed some of recent news stories about Neandertals.  Did they throw spears or javelins?  Does it even matter?  Why would anyone want to hurt Shanidar III?  And do their teeth have anything to say about their love lives?

Julien from A Very Remote Period Indeed discusses the recent paper about Neandertal pigments in context with other Neandertal pigment and shell ornament papers.  A fragmented horse metatarsal?  Very clever.

Michelle wrote us another great post- this one about a possible Homo erectus hearth in Israel.

What if you could go back in time and sit around that ancient hearth?  Martin from Aardvarchaeology discusses what it would be like and why it would be scary.

Magnus from Testimony of the Spade reviews To Wake the Dead, a book about Cyriacus of Ancona and the birth of archaeology.

Julien from A Very Remote Period Indeed relates a call to us paleo-people to integrate all of our cataloged fossils to a better picture of the history of biodiversity.

Another post from Julien discusses some of the recent literature on the archaeology of modern human behavior in East Asia.

At Neuroanthropology, Greg Downey discusses a recent article about the anthropology of American mental illness and other culture-bound syndromes.  What a thought-provoking post!

Over at Anthropology in Practice, Krystal D’Costa was prompted by the Hadza to take stock of her essential items.  My essential item right now is a mug of hot chocolate.

Krystal writes another great post on gold as an internal currency in South Asian culture.

Kerim has a post about becoming expat teacher near the location of his old field site.

Finally, here is one last gratuitous picture of a gelada, because this is my blog, I just submitted a first draft of my thesis, and I do what I want!

Julien will be hosting the next edition of the Four Stone Hearth over at A Very Remote Period Indeed.  Make sure to submit some nice posts to keep us all busy.  I’m looking at YOU, linguists!

So… Did knuckle walking evolve twice?

Almost certainly.

We had lots of clues that this was the case before Ardi, but now that we’ve got Ardi- the palmigrade extraordinaire, we know that humans did not go through a knuckle-walking phase, and that chimpanzee knuckle-walking has evolved since the split with our last common ancestor with them.  Which would also means that it evolved after our split with the gorillas… which means that knuckle-walking evolved twice.

The Great Auk by James Audubon

As we’ve discussed before, knuckle-walking is a pretty weird thing to do, which is why the idea that it evolved only once is hard to shake.  But once you’ve got a particular body plan, there are only so many ways to accomplish a certain task.  For example, the now-extinct Great Auk was a flightless sea bird that hunted fish underwater.  It was white on its front, and black on the back, and had powerful rear feet and webbed toes.  Sound familiar?  The Great Auk was the Northern Hemisphere’s version of the Penguin, but the two were not particularly closely related.  It’s simply that, once you’ve got the body plan of a bird and you want to start diving for fish at high latitudes, you’ve got to rework the wing a little bit so that it’s no longer any good for flying in the air- but man, will it be good for underwater flight!  And then you can work on your body shape a little so that you’re like a little avian torpedo.  And then you’ve got to put some body fat on so that you can withstand the frigid ocean temperatures.  And voila!  You’ve got two almost-identical ocean birds separated by an entire planet and a couple of hundred thousand years, if not more.

The apes have done a very similar thing.  The general Miocene ape was an above-branch quadruped.  In order to get a flexible shoulder, the ape had to move its scapulae so that the glenoid fossa, where the humerus articulates, was facing laterally instead of ventrally.  The glenoid fosssa also became a little shallower, so that it could accommodate a wider range of motion.  When monkeys, or squirrels, or dogs land on the ground with their front feet, the forces are allowed to travel all the way up the humerus until they reach the glenoid fossa.  But think… if you’re an ape with a fancy new flexible shoulder, and you try to land on your front feet, what happens?

Well, you’ll dislocate your shoulder.  That’s a very bad thing for an ape to do!

Monkey Thorax. From Aiello and Dean's Human Evolutionary Anatomy

Ape Thorax (in this case, a human). From Aiello and Dean's Human Evolutionary Anatomy.

Another thing that some of the apes have done is make their elbow more flexible, as well.  If you look at a monkey’s (or squirrel’s, or dog’s) ulna at the elbow, they have a huge chunk of bone on the end of the olecranon process that we don’t have.  Also, the little U (U for ulna!) that cups the trochlea of the humerus points to the side in all of those quadrupeds- more like a C than a U.  But in apes, it points straight up.  All of that makes for an extremely flexible elbow- and gives us the same problem with dislocation that we had with the shoulder.

Olecranon processes (proximal ulna) in a human (left) and a cat (right)

So, you’re a Miocene ape, and you’ve evolved this flexible shoulder and flexible arm, but you’re also starting to get quite large.  So large, in fact, that sometimes you have to leave the trees because the terminal branches that connect one tree to another just aren’t large enough to support you.  So, you come down out of the trees and walk on the ground.  And man, are your muscles working overtime to keep those bones from dislocating!  Each step you talk with your palms on the ground sends a huge amount of force up through your wrist to your elbow, and then up through your shoulder.  Your muscles are concentrically contracting, trying to keep everything in its right place- but the problem with concentric contraction is that you’re already contracting as hard as you can, so one misplaced step, and your muscles can’t do anything to help you avoid injury.

But if you walk on your knuckles, it doesn’t do that!  Because you are such an excellent climber, you have these massively strong flexor tendons in your fingers.  If you flex them and put your weight on your knuckles, those tendons eccentrically contract and are able to absorb some of the force coming at you from the ground.  That way, your arm muscles don’t have to work so hard simply to keep your bones from dislocating.  Plus, you’ve got those long fingers for vertical tree climbing and suspension, and tucking them under your hand gets them out of the way.

Gorillas and chimps have both figured this out (anatomically speaking- I don’t know if any of them have put any thought into it!).  And orangutans are fistwalkers- but I know that I’ve seen one on a really hard concrete surface using his knucles.  It’s a classic case of convergent evolution- but you can’t really tell from the genetics or the living anatomy of the animals- we needed a fossil to finally be relatively certain.  Much like the Auk and the Penguin, the Chimp and the Gorilla faced similar problems brought about by their environment and anatomy, and ended up solving them in similar ways.

Lovejoy, C., Simpson, S., White, T., Asfaw, B., & Suwa, G. (2009). Careful Climbing in the Miocene: The Forelimbs of Ardipithecus ramidus and Humans Are Primitive Science, 326 (5949), 70-70 DOI: 10.1126/science.1175827

Ward, Carol (2007).  Postcranial and Locomotor Adaptations of the Hominoids.  Handbook of Paleoanthropology.  DOI: 10.1007/978-3-540-33761-4

NPR: The invention of osteopenia

All Things Considered ran an interesting story about the birth of osteopenia as a treatable disease.

Bone densitometry becomes increasingly available. And women start wanting it, and they hear their friends have had a measurement of bone density, and their friend was told that they have osteopenia, and they want to know whether they should be treated. And so, it’s almost viral.

I can’t offer any commentary from a medical or clinical perspective, but as an evolutionary anthropologist, I wonder how many of these women could benefit from weight-bearing cardio (walking and running) and strength-building exercises?  As people age, their bone density naturally decreases unless they actively work to maintain it.  It’s a shame that I only very rarely see women over 40 in the weight room doing deadlifts and squats.

The Seed-eaters

If someone stuck a gun to my head and told me to pick my favorite primate, I think I’d have to go with the Gelada (Theropithecus gelada). Just look at him!  Have you ever seen anything more handsome in your whole entire life?

Gelada Yawn

Come on! He’s a freakin’ beatiful monkey, that’s what he is. I dare you to tell him otherwise.

Perhaps my fondness for the most beautiful monkeys on Earth is why Clifford Jolly’s The Seed Eaters has a special place in my heart.  Around the blogosphere, we’ve been talking quite a bit about human origins modeling for the past few months, and while we’ve disagreed about a lot of things, I think the discussion has been interesting and thoughtful.  Jolly’s model has since been falsified, but it shows us that even if a model is incorrect, it can be incorrect in an interesting way.

In The Seed Eaters, Jolly summarized many of the origins models that had been prevalent at the time:  That tool and weapon use led to feminized canines in males, bipedalism, and encephalization; Holloway’s hypothesis that canine monomorphism was the result of selection for hormonal monomorphism and cooperation; that bipedalism was the result of a display posture; and the “Man the Hunter” idea that once humans moved to the savannah, meat-eating and hunting took on special significance and acted as selective forces on bipedalism, and allowed brain size to increase.

Each of the models above tries to answer at least one question about humans:  Why don’t we have any canine dimorphism?  Why are our brains so much larger?  Why are we the only ape that habitually walks upright?

At the time that Jolly wrote his paper, the only fossil humans that we had came from the savannas, so paleoanthropologists figured that one thing that differentiated us from the other apes was that our ancestors had left the wooded forests and lived on the savanna.  We now know that bipedalism evolved before we left the forest, so life on the savanna doesn’t explain very much about our origins.  However, Jolly didn’t know that.

Jolly’s model was a comparative model that made use of geladas, mandrills, and baboons.  Mandrills and many baboons live in wooded environments, but the gelada has become remarkably committed to life in the open grassland.  Jolly asked, What if the primary dietary shift in human origins wasn’t from fruit to meat, but from fruit to grasses and seeds as happened in the gelada?

Jolly made an impressive list of characters shared between humans and geladas but not baboons or chimps, including

• extremely opposable thumbs
• relatively non-opposable great toes
• dimorphism in the hair around the face and neck
• female epigamic features on the front and back
• short and broad cranial bases
• anterior migration of the temporal muscle
• many features of the teeth, such as small incisors, crowded cheek teeth, and early canine eruption relative to the molars

Of these, the ones that I find the most interesting relate to the teeth, because we *do* see parallelisms between geladas and the robust australopithecines in many of these features, particularly in canine size.  Canines and incisors are extremely reduced in the more robust species of human.  Jolly points out that in geladas, the canines have not been reduced because of any behavioral changes, but that their reduction is probably the pleiotropic result of selecting for massive molars.  You’ll still see this argument made for the reduction of canine size in humans.  However, with Ardipithecus, we see that the canine reduction is not accompanied by an increase in molar size, so it probably cannot account for the initial shift.

(Aside- I know from the picture above it doesn’t look like the gelada has reduced or monomorphic canines at all!  But compared to fossil Theropithecus species, they actually are.  In male geladas, the largest individuals have the smallest relative canines.)

Jolly also points out that, in many of the savanna-dwelling species, the typical one-male/multi-female group lives within a larger “troop” composed of many of those one-male groups.  The males within the troop cooperate with each other when defending their groups against extra-troop males.  This idea is relevant to the recent discussions of Ardi as well.

Jolly concludes his paper by outlining what he thinks a good origins model should be:  It should have testable predictions (which can be falsified by new fossils), and it should be able to account for data from different fields.  It should not only be plausible, but convincing.  As I said above, Jolly’s model fails now based on his own criterion of falsifiability, but I think that’s what makes it stand out in my mind.  Many of the origins models that we discuss today have no way to be falsified, so they stick around because they are plausible.

The Seed-Eaters: A New Model of Hominid Differentiation Based on a Baboon Analogy, by Clifford J. Jolly

© 1970

Wednesday Morning Giggle

This is the first dinosaur bone ever described.  It hails from Oxfordshire, England, and was discovered in 1676.  It is the distal femur of a very large animal which we now call Megalosaurus.

When it was first described by Oxford University chemistry professor Robert Plot, he knew that it was a femur, but decided that it belonged to a race of Antediluvian giant humans.  It was far too large to belong to anything that was alive at the time, and his guess was perhaps the most likely possibility based on the knowledge of the time.

In 1763, Richard Brookes got ahold of the bone and recognized that it was not that of a human, giant or otherwise.  He named the new creature Scrotum humanum, and his description relayed the uncanny resemblance of the bone not to the distal femur of a human, but to another one of the human’s anatomical features.

For whatever reason, the name didn’t stick.   Thankfully, we do not have an entire genus of dinosaurs with the name Scrotum.  We also no longer believe the bone belonged to a Pre-Noachian giant human.

I find these stories of mistaken fossil identity fascinating.  Homo diluvii testis turned out to be a giant salamander, and Scrotum humanum was a giant dinosaur.  The world looks so different to us now then it did back in the 18th Century, when giant humans and worldwide floods were still on the cutting edge of science.  We’ve come a long way, baby.

Repost: The General Ape Body Plan

All of the extant apes are adapted to suspensory, upright locomotion.  Their forearms and hands are elongated in order to reach different branches without leaping.  The glenoid fossa in their shoulder faces laterally and is shallow, allowing them to circumduct their arms.

The large-bodied African apes have a shortened spine compared to other primate species.  This is particularly evident in the lumbar spine.  Primates typically have 7 lumbar vertebra.  The lesser apes have 5-6, and great apes have 3-4.  This reduction in lumbar vertebra number lowers the rib cage, and the iliac blades of the pelvis have extended cranially.   The rib cage is in roughly the same plane as the top of the pelvis, which offers a stronger base for muscles like latissimus dorsi to act on the upper limb.  This “short-backed” body form also serves to protect the spine against lateral bending moments when latissimus dorsi contracts.  This protection is provided by the erector spinnae group in most other quadrupeds, but in order to get the glenoid fossa so that it faces laterally instead of ventrally, the spine migrated into the thorax, and the erector spinnae was reduced.

Thoraxes drawn by Adolph Schultz

Apes also share a feature called “ulnar withdrawal,” which allows them the ability to adduct the wrist.  The styloid process of the ulna is reduced and is not in contact with the bones of the carpus (namely, the triquetrum and pisiform).  A small bone called the pisiform has moved distally.  Where once there was a “wrist mortise” achieved by the ulnar styloid and the carpal bones, there is now a semilunar mensicus.

The fingers of the apes have become elongated, except for the thumb.  These long fingers are probably an adaptation to both suspension, in which the ape hangs from a brach, and vertical climbing, in which the ape wraps its hands around a vertical support.  The apes use their fingers as a “hook” when they hang from branches, and this requires strong musculature in the hands.  Their halluces associated with very strong musculature as well, reflected in a well-developed fibula.

Gibbons are highly derived apes, both in body size and in their locomotion.  They are so good at brachiating that they seem to ricochet from one branch to the next.  Gibbons have evolved extremely elongated limbs and fingers, but haven’t done much with their thumbs at all.

Orangutans almost never come down to the ground, but instead move around the upper canopy of the forest.  Their very large body size requires them to be very careful, and their locomotion is very deliberate.  They use both hands and feet with equal proficiency, and have very flexible joints as a result.

A chimp places its weight on the middle phalanx.

Chimpanzees are arboreal for much of their time, but also spend time on the ground as knuckle-walkers.  Gorillas share both of these modes of locomotion, but spend much more time terrestrially as a result of their extremely large body size.  Knuckle-walking is an adaptation which serves as a compromise between arboreal suspension, which requires long hands and fingers, and quadrupedalism, during which long fingers can be cumbersome.  In addition, knuckle-walking allows the strong digital flexor muscles and tendons to absorb some of the forces from the ground before they reach the flexible joints of the upper arm.

It is helpful to remember that even though extant apes share a certain suite of features, each has their own special adaptations to their own environment.  Some of these anatomical characters are shared as the result of common ancestry, but some may be convergences resulting from similar selective pressures.  This is particularly important to remember when reconstructing the fossil record of these apes.

Miocene “Monkey”: Pliopithecus canmatensis

What could possibly be a better Christmas present than a new fossil primate?  Nothing, that’s what!

The most recent addition to our family bush is a Pliopithecine from Spain named Pliopithecus canmatensis.  Pliopithecoids are gibbon-like in many ways, including their long limbs, large hands, and maybe the ability to brachiate.  However, the pliopithecoids are much too ancient to be directly related to gibbons, and probably predate the split between monkeys and apes. Resemblance to extant gibbons is almost certainly an example of convergence.  Pliopithecoids have two premolars, which connects them with the catarrhines, but they also have unique dental morphology which places them in their own group.

The scientists describing the fossils- David Alba and his colleagues- have outlined an evolutionary history for the pliopithecoids in which they were the first catarrhines to leave Africa for Eurasia.  In the Early Miocene of Asia, these first pliopithecoids are represented by a group discovered in 1978 called the dionysopithecines.  From this group evolved the true pliopithecines.  By the Middle Miocene, pliopithecines had dispersed quite happily into Europe, and then by the Late Miocene, back into Asia.

P. canmatensis is your typical toothsome primate:  8 individuals are represented from 61 teeth and a few scraps of mandible and maxilla.  From the maxillae, we can tell that the face was short.  The madible is long and skinny, with tooth rows that are almost parallel to each other.

There is only one upper canine tooth represented in the sample, and it is probably from a female.  It has wear facets that show that it occluded with Premolar 3 and the lower canine.   There are a few lower canines, and they probably display sexual dimorphism in size and shape.  The female canines have a blunt surface, while the males have a pointed apex.  The female’s canines are gracile, while those from males are “stout,” but taller than the female’s. Premolar 3 is a typical honing premolar:  It only has one, high cusp with a distinct, steep wear facet which resulted from consistent contact with the upper canine.

In the molars, the distinctive “pliopithecine triangle” is present in the upper 2nd and 3rd molars.  This triangle lies on the cheek side of the tooth, in between the protocone and the hypocone, and even with the beautiful diagram which the authors provide, I have to squint my eyes and cock my head to see what could maybe pass for a triangular shape of some sort.  But that’s what I have to do for all teeth, so I trust the authors that it’s there!

The sites at which these fossils were found-  the Hostalets de Pierola-  are becoming quite the treasure trove of primate fossils!  The area in Catalonia is the home of of the most interesting Miocene hominoids, Pieorolapithecus catalaunicus.  Anoiapithecus brevirostrus and a speices of Dryopithecus also hail from the area around Barcelona.

Alba, D., Moyà-Solà, S., Malgosa, A., Casanovas-Vilar, I., Robles, J., Almécija, S., Galindo, J., Rotgers, C., & Mengual, J. (2009). A new species of Gervais, 1849 (Primates: Pliopithecidae) from the Middle Miocene (MN8) of Abocador de Can Mata (els Hostalets de Pierola, Catalonia, Spain)
American Journal of Physical Anthropology DOI: 10.1002/ajpa.21114

An interesting thought

“… it is illogical to invoke the behaviour of living apes to explain the origins of something that they themselves have not developed…”

The Seed-Eaters: A New Model of Hominid Differentiation Based on a Baboon Analogy, by Clifford J. Jolly © 1970