I'm here on the Sandwalk, where Darwin walked almost every day to think about his deepest problems,

and, maybe the deepest of all, was the nature of species. Why are species bounded from each other? Many people object it. If Darwin's ideas about gradual evolution over time were correct, then we ought to find, not only, many, many gradual changes in the fossil record, we should also find gradual gradations among living organisms. You shouldn't see that there are distinct types from place to place, you should see a continuous range. In fact, Darwin pointed out, in the Origin of Species, that this was, in his mind, one of the most fatal problems that he faced. He had an explanation. Darwin's view was that as gradual changes occurred within species, and they became more and more adapted to small habitats, those gradations would give rise to larger populations in some places and naturally smaller populations in very narrow habitats. In Darwin's point of view, the variation within larger species would be continually giving rise to more and more favorable variations, so that the larger species would continually replace the smaller ones. This idea, the idea that populations that were large were ultimately population sources and smaller, more narrowly adapted species were always vulnerable to incursions from these larger groups, that idea is still fundamental to our ideas to the nature of species and their evolution over time. Of course, in Darwin's day,

the fossil record was not very complete. Today, we have extraordinary records of the evolution of some lineages. And the hominins are one of them. One of the most important examples of gradual change over time, in a single geographic region, is the evolutionary trajectory from Australopithecus anamensis to Australopithecus afarensis. These are two species that lived between about 4.2 and about 2.9 million years ago across much of East Africa. And when we look at these species and the fossil record that preserves them, what we see is a record of impressive, gradual change. Today, biologists use different concepts of species to apply to different groups of animals. And in particular, in hominin evolution, we use a different concept of species to talk about living animals and living primates than we do to talk about fossil primates and fossil hominins. That can cause a lot of confusion. Now in Darwin's time, species were basically what naturalist's said they were. Naturalists went out, they began to understand the variation, they understood, in particular, the morphological variation, and they gave animals names based on their understanding of that variation. I've taken some time to sit down on the bench here at the end of the sand walk, and I'm reading through a copy of, The Descent of Man. This is Darwin's book about human origins, where he talks about human variation, and he's concerned with the topic of whether different kinds of humans should be called

different species or not. And he addresses this question in a way that really tells us that Darwin was grappling with the problem of species just as today's paleoanthropologists do. He wrote, "It is a hopeless endeavor to decide this point," whether different groups of humans should be called different species, "until some definition of the term species is generally accepted. We might as well attempt, without any definition, to decide whether a certain number of houses should be called a village, town, or city. We have a practical illustration of the difficulty in the never-ending doubts whether many closely allied mammals, birds, insects, and plants, which represent each other respectively in North America and Europe, should be ranked as species or geographical races." When paleoanthropologists are naming species in the fossil record, we're naming them for reasons that sometimes have little to do with whether we really think these were different, biologically-separated groups or not. We want ways to talk about them. We're using different concepts of species. The use of different concepts of species, by biologists for different purposes, today may be more sophisticated in the way we talk about it than in Darwin's time. But the problem is often fundamentally the same. The evidence that we use to decide whether species have a close evolutionary relationship is not often evidence that's closely at hand. Most biologists today, when they talk about living mammals, in particular, living primates, use a concept

of species called the biological species concept. Under the biological concept of species, a species is a population that includes all of the populations that can actually, or potentially, reproduce with each other in their natural habitat. The biological concept of species focuses on reproduction. It doesn't, in principle, matter how different these organisms look, as long as they're reproducing with each other in their natural habitats, in their natural populations. We have much more evidence about the nature of variation among populations of great apes. In particular, when we look at chimpanzees, which exist today in at least four different subpopulations across equatorial Africa, their close relatives, the bonobos, who separated from the chimpanzees more than a million years ago, we can really begin to look at the historical importance of gene flow between these subspecies. Chimpanzee subspecies have existed for more than a half million years. And that variation, it turns out, is very comparable to the variation that we're going to see in the later hominin fossil record, when we look at populations like the Neanderthals. But when we look at early hominins, we don't have very much evidence about whether those early hominin populations were comparable to subspecies, as we see in chimpanzees, as we see in gorillas today and orangutans, or whether that variation is comparable to species, as we see between bonobos and chimpanzees.

In some ways, the difference between subspecies and species is sort of arbitrary. Today, many conservation biologists, who are concerned about protecting chimpanzees, really want to emphasize the biological differences between groups, and would prefer that the different subspecies be called different species. Likewise, many of us who study the fossil record, who are trying to figure out how these different fossil forms, who are morphologically different, might be related to each other., want to emphasize the morphological characteristics by giving things different names, and aren't yet able to test whether those morphological differences might be consistent with populations that are exchanging a lot of genes. So the Australopithecus afarensis is the descendant of Australopithecus anamensis, and it looks like anamensis evolved gradually and afarensis is a continuation of that same evolutionary trend, why don't we just call them one species? Why isn't it the one evolving species of Australopithecus afarensis? Here's why. Even though the fossil record of hominins is one of the more complete for any lineage that we have, it's still missing a lot. We know what existed in East Africa, from Ethiopia to Tanzania, around the time that these species existed. But, we don't know everything that existed then. And, even though afarensis might be the descendent anamensis, we don't know that it's the only descendant of anamensis. Hominins were diversifying. In the slightly later hominin fossil record, we have evidence for many different kinds of

hominins that are adapting to different kinds of ecologies. We have hints across Africa in the early to middle [inaudible], the time that anamensis and afarensis are evolving, we have hints of diversity. There is, from Chad, a specimen from a site called Bahr el Gazel that's been tentatively assigned to the species Australopithecus bahrelghazali. We have a group of fossils, including one relatively complete skull from Kenya, that have been assigned to another genus, Kenyanthropus. Kenyanthropus platyops. Those things might represent different descendants of anamensis, or they might represent separate lineages that have common ancestors with other hominins before anamensis. We just don't have enough evidence about them to say. And so, until that time, paleontologists prefer to give things names so that they can communicate about them clearly. We know that we can document gradual evolution in the hominins, but what we don't know is exactly when and where major changes might have arisen. The bottom line is, biologists use different concepts of species for different reasons. When we look at the fossil record, some of the species that we talk about are species that are really well-documented by large fossil samples where we can begin to test the idea that they could, potentially, reproduce with each other. In other cases, we name samples, which might consist of a single, individual specimen, because we want to talk about them clearly. We may not be able to test whether those samples belonged to common populations with things that are morphologically different, but the naming gives us,

scientists, a way to deal with that variation and to approach the hypotheses. When we argue about whether a species is valid or not, we're not arguing about whether evolution happened, what we're arguing about is whether we can trace the pathway of evolution with the fossil record that have or whether we need to make additional discoveries to find the common ancestors of these groups. I hope you've enjoyed this visit to Darwin's Sandwalk to talk about the nature of hominin species. It is a terrible problem for us today as we try to teach the fossil record, because there are so many names. But when you understand how we're using those names, you can begin to focus in on the important differences between samples of fossils, and our ability to test hypotheses about them. It is a very good book.