How many apes do you know? I don’t mean as individuals but as types, or if you wish to be scientific about it, as genera: subdivisions of biological families. Well, I’m sure you can name the chimpanzee (which has the generic name Pan) and the gorilla (Gorilla) for the African apes, and they are indeed the only two African genera that exist, with three or four species within each. I suspect you will name the orangutan (Pongo, two species) and probably the gibbon (Hylobates, twelve species) as Asian apes.
I would certainly hope you appreciate our evolutionary heritage and list humans (Homo, one species) as another type of ape, but I would be concerned if you mentioned baboons, mandrills, capuchins, marmosets, or howlers, all of which are types of monkeys and quite different from apes. I’d be impressed if you named the siamang, which is in the same family as the gibbon, the Hylobatidae, otherwise known as the lesser apes, and provides a further three genera (Nomascus, Hoolock, and Symphalangus). You couldn’t get any further, at least with apes alive today, because there simply aren’t any more to name. But you’ll probably be familiar with Australopithecus as an extinct ape genus to which Lucy, Australopithecus afarensis, belonged some 3.5 million years ago (MYA). And if you follow the human evolution literature you might even cite Ardipithecus from 4.4 MYA.
Forgetting about the siamangs, the other seven types of ape—Homo, Pan, Gorilla, Pongo, Hylobates, Australopithecus, and Ardipthecus—were all that I could have confidently mentioned before becoming familiar with The Real Planet of the Apes. I might have stumbled over a couple of other challenging names, Ramapithecus and Sivapithecus, vaguely recalled from student lectures in the early 1980s about their controversial part in human evolution. But within the pages of David Begun’s book, primarily concerned with the Miocene world between 23 and 5 MYA, I’ve now encountered no fewer than fifty genera of apes, ranging from Aegyptopithecus at 33 MYA from Egypt to Indopithecus at 6 MYA from Pakistan, and eventually Gigantopithecus, an ape the size of a polar bear that dwelt in the forests of Southeast Asia until a mere 300,000 years ago. And that is probably just half the number of ape genera that have existed on earth.
I trust that David Begun, professor of anthropology at the University of Toronto, could have reeled off all fifty tongue-twisting pithecus names without any deviation, hesitation, or repetition. He is a world-leading authority on ape evolution and has gotten to know many of them as intimately as one can by literally pulling them out of the ground, poking about in their nasal cavities, and meticulously measuring their finger bones to write what has been (lovingly) described as one of the most boring academic papers of all time. His new book is enthralling, making accessible an absolutely critical period of human evolution and the scientific debates surrounding the interpretation of its evidence—including his own controversial thesis.
The last two decades have seen a plethora of books about human evolution, many written by leading anthropologists and often quite superb at expounding the last six million years of evolution, taking the common human-chimpanzee ancestor as their starting point. They typically move swiftly to the appearance of the Homo genus in Africa with its Oldowan stone tool technology at two million years ago. They continue with the study of bipedalism, brain enlargement, extended life history (notably childhood), and Old World dispersal. They explain the diversification into multiple species of the Homo genus, including H. ergaster, erectus, heidelbegernsis, and neanderthalensis, and how advances in evolutionary genetics are helping us to understand the relationships between the most recent of these species. As indeed are new finds, as the discovery in 2013 of Homo naledi testifies (it was announced to the world in September 2015). Hence these books will keep flowing. Typically, they end with the evolution of symbolic thought and language, either as a slow burn or a transformational event. Homo sapiens is left alone as the only living member of our genus, which is gradually pushing the few other remaining apes into extinction by its reckless behavior.
These accounts are fine but increasingly seem inadequate, or at least only part of the story. The more we learn about our ape cousins—their tool-making, social intelligence, communication skills, complex social behavior, and cultures—the more it appears that much of natural selection’s work for human evolution had been undertaken long before the time of not only the common human-chimpanzee ancestor but also those ancestors we shared with the gorilla at about 10 MYA and the orangutan at about 15 MYA. While the brain of the chimpanzee and the australopithecines may be three times smaller than our own, their brains are nevertheless more than three times larger than what is expected for a mammal of their body size. So an evolutionary trajectory had been set many millions of years before Homo emerged.
For a complete story of human origins we must look back in time to at least thirty million years ago. Begun begins his account at Fayum, now located in the desert of northern Egypt but then amid a coastal subtropical forest. It contained an astonishing assortment of animals, some that would appear familiar to us and some quite strange, including elephants the size of cows, crocodiles, turtles, and whales with arms and legs. In the trees sat Aegyptopithecus, looking like a cross between a monkey and a lemur, picking and peeling ripe fruit. This primate, known from an assortment of fragmentary fossils, is likely to resemble the common ancestor between Old World monkeys and apes, their evolutionary divergence being dated by the molecular genetic clock at thirty- one to thirty-eight million years ago.
That does indeed mark an appropriate starting point. Although monkeys are certainly both clever and adaptable, apes—at least those alive today—are an order of magnitude more intelligent. Apes live longer. They have more complex societies and bigger brains. They provide an irresistible comparison to humans, having the same body plan of straight backs and long arms that evolved for “suspensory” movement—hanging from branches. They alone have a female menstrual cycle (monkeys have a seasonal reproductive cycle) and habits of using tools and building nests—ours being far more elaborate in scale and style, but in principle just the same. We are indeed merely a type of ape, the only one that uses bipedal walking as its primary locomotion rather than swinging from branch to branch, leaving monkeys to walk on all fours along the tops of branches.
Fayum at 33 MYA provides the start and Ethiopia at 4.4 MYA the end of Begun’s account, this taking us to Ardipithecus, which lived immediately prior to the spread of the australopithecines, which is when most accounts of human evolution begin. To understand what happened between these two African localities we have to travel via Europe because that is where, Begun proposes, much of the evolutionary action occurred.
This will sound like heresy to some. After an undistinguished and quite frankly racist history of Eurocentric research throughout much of the late nineteenth and early twentieth centuries, we have all become at ease with being “Africans.” Hasn’t the fossil, archaeological, and genetic evidence convincingly demonstrated that Charles Darwin was correct when he proposed in his Descent of Man (1871) that “it is somewhat more probable that our early progenitors lived on the African continent than elsewhere”?
Well, yes, with regard to Homo sapiens, for we all descend from populations that dispersed from Africa a mere 70,000 years ago—with a tiny contribution from a few Neanderthals and others. But where did those early African progenitors come from? From European apes, Begun proposes, apes that dispersed into Africa around ten million years ago already having the relatively big brains and extended life histories that, after further elaboration, would become a distinguishing feature of the Homo genus. So while Africa may have been the cradle of humankind, Europe had been its birthing pool.
“Europe,” of course, is a problematic term for a northern segment of the Miocene world, which existed between circa 23 MYA and circa 5 MYA. This was the segment that was undergoing transformation by tectonically driven mountain building that created the Alps, Pyrenees, and Zagros. As with “Africa,” we cannot read the word “Europe” without thinking of the cultural construct that this term represents today and the history of its relations with Africa. When the evolutionary story includes accounts of “migrations” and “dispersals,” there is a resonance with events happening today as refugees flee into Europe from Syria and North Africa that is almost overwhelming, but, of course, hugely misleading. For here we are dealing with evolutionary time, millions of years, with the fossil record providing no more than tiny and very occasional glimpses of what was happening within the Miocene world.
Before “Europe” became center stage, the apes themselves had to reach that northern landmass. For the first 16 million years they remained entirely in Africa and provide a fascinating cast of characters within an evolutionary story of diverse adaptations. I say “characters,” but they all have no more than bit parts, some of which are very bitty indeed. Kamoyapithecus, for instance, found in Kenya and dating to 26 MYA, is known only from a fragment of upper jaw and a few isolated teeth, while Rukwapithecus, from the same locality, has (so far) left us with no more than a lower jaw. Teeth and jaws are indeed the most prominent finds because of their durability, with the tiny humps and bumps on the molars and the dental characteristics—the number of incisors, canines, premolars, and molars—being absolutely critical to identifying the dietary behavior and evolutionary affiliation of the find.
Paleoanthropologists have the enormous challenge of using such dental minutiae to reconstruct the evolutionary relationships between one fossilized jaw and another, separated by millions of years. When bits of limb bones are found, proposals can be made for posture and mobility, the shift to an upright posture and brachiation—swinging from branches—being critical evolutionary transitions. Paleoanthropological heaven is when jaw and limb fragments are found together. This is the case for Morotopithecus from Uganda, dating to around 20 MYA, which generously left behind a jaw, a couple of vertebrae, a shoulder blade, and a hip joint. When put together they create a curious ape that appears dentally primitive but advanced in posture, indeed not very different from the way modern apes move around in trees. The dilemma we face, however, is whether the bits and pieces should actually go together at all. They might derive from several different types of apes from different time periods that have become aggregated into the same geological deposit, leading paleoanthropologists to create no more than an imaginary Miocene chimera.
I don’t blame them for a few accidents like that. Begun is refreshingly open about the speculation and imagination sometimes required within his science, and the mistakes that have been made. Another unavoidable but irritating habit of paleoanthropologists is that they change their minds about what genus a fossil belongs to or what it should be called. One of the earliest African ape fossil discoveries (1933) was named Proconsul (a name unfortunately inspired by a popular performing chimpanzee of the time called Consul) dating to 19 mya. Some specimens of Proconsul have now been renamed as Ekembo. Far from being a bit player, Ekembo is known from almost every bone in its body and evidently looked and behaved much like a monkey—with the exception of lacking a tail. This lack of a tail is a distinguishing sign of an ape, or more accurately one should refer to the presence of a tailbone, the coccyx. Quite why tails were dispensed with is unknown but Begun manages to weave a marvelous evolutionary story that connects the loss of the tail to the evolution of spoken language, which is speculation and imagination being put to very good use.
This appearance of a new type of ape, Afropithecus, at 17.5 mya, brings to its climax what Begun calls the first “golden age” of apes or near apes of the African early Miocene. The hugely fortunate find of a baby’s jaw enabled the age at which Afropithecus’s first molar erupted to be estimated. This is probably the most important measurement that can be made because it correlates with a range of attributes that are otherwise impossible to observe, including life span, age at menarche, lifetime number of offspring, and brain size. While the age of eruption of the molar in Proconsul has been estimated to be the same as in modern-day monkeys, that of Afropithecus was found to be intermediate between monkeys and apes, suggesting a relatively larger brain size and a longer period of infant dependency and learning.
That innovation in life history together with new dietary adaptations might have been critical for the dispersal of apes from Africa into Eurasia, occurring soon after the appearance of Afropithecus. The route taken was from East Africa and Saudi Arabia, this being made possible by the breakup of the vast early-Miocene Tethys Sea that had stretched from the Atlantic to the Indian Ocean. As this broke up, land bridges were created, utilized not just by apes but by a host of other land mammals to disperse north from Africa.
The first apes in Europe are grouped together as griphopiths, described by Begun as the first “hominids”—the group that today contains only the great apes: Pongo, Gorilla, Pan, Australopithecus, Ardipithecus, and Homo. This is to be distinguished from the “hominoids,” encompassing all the apes (including the gibbons and siamangs) and the “hominines,” which contains just the African apes and their descendants (and hence humans). Finding the first hominid fossil sounds momentous and Begun provides an evocative description of what its location at Engelswies in Germany would have been like 17 million years ago—but then we learn that the evidence consists of no more than just one half of a tooth, and a worn one at that. The fossil record is better at Paşalar in Turkey, showing that the griphopiths had especially robust jaws and thickly enameled teeth. Another form of ape is also found at Paşalar: Kenyapithecus. As the name implies, this has also been identified in Africa, suggesting a series of ape expansions between Africa and Eurasia during the Middle Miocene.
The traffic was both ways, with griphopiths spreading into Africa and diversifying into a number of new genera. The best known is Nacholapithecus, from Kenya at 15 mya, which left us the most complete fossil of an ape from the Miocene—and hence no risk of mixing up the body parts from different creatures. Nacholapithecus had a strange combination of anatomical features, unlike any living primate, with a trunk like a monkey, a coccyx (hence it was an ape), and forelimbs that suggest a specialized grasping capacity even though it did not swing from branches. This exemplifies the delight of the fossil record, showing us the reality of creatures that we might only otherwise imagine in a childlike manner, evolution constantly experimenting to fill adaptive niches with animals that ultimately had no future in the world.
After 13 million years ago the fossil record in Africa dries up while that in Europe and Asia does the opposite. It not only increases but shows the development of local apes, most likely reflecting restrictions on dispersals and hence gene flows between and within Africa, Asia, and Europe arising from the ongoing emergence of mountains. New ecological pressures stimulating natural selection were created by the constantly changing topography of valleys, highlands, and coastlines. Indeed around 12 million years ago a fundamental divergence in ape evolution occurred, creating two lineages: the Asian pongines, from which the present-day orangutan derives, and the African hominines, although the latter were at this time located in Europe, not Africa.
The Asian evidence is especially complex and has suffered from what Begun describes as a spectacular amount of taxonomic shuffling as paleoanthropologists changed their minds about species and genus names. The most sorry—but fascinating—saga concerns Ramapithecus and Sivapithecus—the controversy I had vaguely recalled when first picking up Begun’s book. You’ll have to read it to understand that story but it captures many of the challenges bravely faced by paleoanthropologists that pervade the interpretation of almost every fossil they find. Are two fossils of the same geological age and locality different in their dentition and brain size because they are from different species, or because they are the male and female or the juvenile and adult of the same species? Conversely, have the similarities in fossils from different time periods and/or localities been inherited from a common ancestor, which would be informative about their evolutionary relationship, or have they arisen by parallel evolution, the two species adapting to similar ecological pressures in similar ways but in complete isolation from each other?
Even before these challenges arise there are those concerning what the specimen looked like prior to the deformation and fragmentation during the process of fossilization. Begun provides examples of when he has taken fossils apart and glued them together again to correct past mistakes. Then there is the question of dating the fossils, especially those recovered prior to today’s meticulous documentation of the locations of discoveries and geological deposits. And then there are three challenges epitomized by the Ramapithecus–Sivapithecus saga: the temptation to “big-up” one’s find as being transformative to our understanding of human evolution (e.g., as Louis Leakey did in 1967) and the likelihood that new fossil discoveries will prove you wrong. While we all pay lip service to the desirability of falsification, we hate it when it happens and often try to deny it. The third challenge is that molecular biology is ultimately much better at deriving evolutionary relationships than paleoanthropologists can ever be at measuring the humps and bumps on fossils.
Back to the evolutionary story. As the Asian apes were evolving toward the orangutan and the now extinct Gigantopithecus, strangely known only from giant molars—its limb bones possibly are sitting in museum basements misclassified as those of a bear—the European apes were evolving along a different track. Dryopithecus, the “oak-forest ape,” was one of the first discovered in 1856, just three years before the publication of Darwin’s Origin of Species, as a collection of fossil fragments at a spot in the foothills of the Pyrenees. The eminent French paleontologist and archaeologist Édouard Lartet interpreted the fragments as deriving from a chimpanzee-like creature, which we now date to around 12.5 million years ago.
Darwin was aware of the discovery and followed his much-cited statement about African origins in The Descent of Man by noting Dryopithecus and the possibility of ape migration from Europe into Africa. That is manna for Begun—Darwin being everyone’s ultimate authority to support a theory. With a considerably larger number of fossil fragments, he now describes Dryopithecus as having been a combination of an orangutan, as identified by its skill in the trees, a chimpanzee, as identified by its diet, and a gorilla, according to the anatomy of its face. It was another of natural selection’s intriguing experiments.
It was from Dryopithecus that genera of apes evolved in Europe with distinctively African ape characteristics. By ten million years ago Hispanopithecus was swinging from branches in the subtropical forests of what is now Spain, its fossilized limb bones indicating that it moved around in a manner similar to modern-day apes, while fragments of crania indicate African-looking sinuses and brow ridges. At a similar date Rudapithecus was hanging out in Hungary, as is evident from its limb bones, digits, and fossilized inner ear that indicates the well-developed sense of balance required by suspensory primates. Rudapithecus also left behind a relatively well-preserved cranium with most of the face and brain case attached, a fossil known as Gabi—paleoanthropologists like to give their finds nicknames. Gabi is the earliest evidence for a chimpanzee-sized brain in the ape fossil record, suggesting that this brain expansion occurred in Miocene Europe.
It is the descendants of apes such as Hispanopithecus and Rudapithecus that—Begun argues—dispersed into Africa around nine million years ago, providing the evolutionary foundation for the present-day African apes and the species that became the global ape, Homo sapiens. Why did they go to Africa? Because beginning fifteen million years ago Europe had been becoming cooler, drier, and more seasonal, conditions unsuitable for apes but ideal for monkeys. So the apes moved out, heading south into Africa, and a population of monkeys moved into Europe.
Critical to Begun’s thesis is that between 12.5 and 9.5 million years ago Africa was effectively devoid of apes. The fossil record is indeed remarkably sparse, leaving an evolutionary gap between the relatively abundant early Miocene apes and those that evolved after 7 million years ago and that are evidently on the course to Australopithecines and Homo. Most notable are Sahelanthropus tchadensis at 7–6 MYA and Ardipithecus at 4.4 MYA, the latter showing another peculiar mix of anatomical traits, defying the idea that there was a linear, ladder-like trajectory toward the evolution of Homo. Had there not been this 12.5–9-million-year gap in the African record, the African ape-like characteristics of the European genera might have been explained as an independent evolution, parallel but not ancestral to the African apes.
Begun’s book reminds us of the hugely impoverished world we live in today, with no more than four types of apes other than ourselves left on earth. Here we must acknowledge that one of those apes—Homo sapiens—is indeed a rather special type. Begun notes how curious it is that gorillas and chimpanzees appear so similar to our early ancestors while humans are so radically different, despite all having had the same amount of time to evolve. Moreover, within the Homo genus, our species appears so radically different to all others, especially Homo neanderthalensis that became extinct a mere 30,000 years ago and had a brain size equivalent to our own.
This is, I suspect, because there was a unique evolutionary development a mere 100,000 years ago within a single lineage of apes somewhere in Africa: the evolution of language. This was not merely a more complex form of the vocal and gestural communication that all apes possess, one that reached its apogee with the Neanderthals. Language was the vehicle for a new type of thought that provided Homo sapiens with their competitive edge over all other species as they dispersed from Africa 70,000 years ago. The use of language created a new dynamic of culture change. In a tiny, very recent moment of evolutionary time, Homo sapiens, the only language-using ape, has created remarkable works of art and scientific achievement, while also contributing to the severe impoverishment of the natural world.
If only Oreopithecus—the last native ape in Europe—could have (literally) hung on for another six million years so we could have seen this wonderful creature that had evolved in a Tuscan swamp, becoming very large but with a small brain, a tree sloth–like ape. I also sensed from Begun’s book that paleoanthroplogy has itself become a little impoverished, and I worry for its future, just as I worry for the gorilla. Not because of the state of relevant science but because of the loss of a type of scientist, some of whom we glimpse in Begun’s account. These are the natural historians who had encyclopedic knowledge of the fossil record of their region and boundless generosity to younger researchers, the likes of Professor Miklós Kretzoi (1907–2005), the Hungarian paleontologist known affectionately to all as Uncle Miklós, and Professor Miquel Crusafonet-Pairó (1910–1983), the Catalan paleontologist always impeccably dressed as a gentleman.
One both hopes but also rather dreads that the ongoing molecular biology revolution will one day extract DNA from the fossils of Aegyptopithecus and Dryopithecus and from Hispanopithecus and Ardipithecus. That will subvert the need for the speculation and imagination required to interpret the meaning of tiny humps and bumps on fossilized teeth and bones for our evolutionary past. If such a scientific juggernaut does ever emerge in the Miocene world—and it may be a road too far—then I hope it verifies Begun’s thesis. For this is not just about a European origin for African apes, because the European apes themselves derived from Africa. Our human genome has been shaped by population dispersals back and forth between the continents, as has the whole history of human culture. That seems a good lesson to remember as hundreds of thousands flood into Europe today.