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Major transitions in human evolution

Robert A. Foley1, Lawrence Martin2, Marta Mirazón Lahr1
[Link] and Chris Stringer3
1
Leverhulme Centre for Human Evolutionary Studies, University of Cambridge, Cambridge CB2 1QH, UK
2
Turkana Basin Institute, Stony Brook University, Stony Brook, NY 11794-4364, USA
3
Natural History Museum, London SW7 5BD, UK

Introduction Evolutionary problems are often considered in terms of ‘origins’, and

research in human evolution seen as a search for human origins. However,
Cite this article: Foley RA, Martin L, Mirazón
evolution, including human evolution, is a process of transitions from one
Lahr M, Stringer C. 2016 Major transitions in state to another, and so questions are best put in terms of understanding
human evolution. Phil. Trans. R. Soc. B 371: the nature of those transitions. This paper discusses how the contributions
20150229. to the themed issue ‘Major transitions in human evolution’ throw light on
[Link] the pattern of change in hominin evolution. Four questions are addressed:
(1) Is there a major divide between early (australopithecine) and later
(Homo) evolution? (2) Does the pattern of change fit a model of short trans-
Accepted: 20 April 2016 formations, or gradual evolution? (3) Why is the role of Africa so prominent?
(4) How are different aspects of adaptation—genes, phenotypes and
One contribution of 17 to a discussion meeting behaviour—integrated across the transitions? The importance of develop-
issue ‘Major transitions in human evolution’. ing technologies and approaches and the enduring role of fieldwork
are emphasized.
This article is part of the themed issue ‘Major transitions in human
Subject Areas: evolution’.
evolution

Keywords:
human evolution, major transitions in human 1. From origins to transitions
evolution, early hominins, evolution of Homo The word probably most associated with our evolutionary past is ‘origins’. The
history of science is awash with books and papers in search of human origins,
Author for correspondence: or the origins of the things that made us human—upright walking or language
or culture. Seeking origins is looking for the beginnings of something, finding
Robert A. Foley
out why and when something that did not exist before did so afterwards.
e-mail: raf10@[Link] Origins research is at its most ultimate in cosmology, when, to the layman at
least, the origin of the universe is when something (matter) is there when pre-
viously (if one can use that word given that time itself did not exist!) there had
been nothing.
Origins research has often been criticized, on both theoretical and practical
terms. Theoretically, the argument has been made that a focus on origins prior-
itizes particular periods and features, and creates essential traits and moments
of significance in a continuity of process [1]. Pragmatically, the search for ori-
gins is a recipe for frustration. There may be a hypothetical point of origin
for Homo sapiens, but to find one fossil closer to that elusive point than another
is only to engender the search for another that is closer still, until the path leads
inexorably to the origins of something else.
And yet we know that there was a time when something did not exist—
humans—and then a time when they did. How do we discover the process,
timing and causes of such a change? This, of course, is not exclusive to
humans, but would apply equally to dinosaurs, mammals, primates and the
most insignificant house louse. How do we square the circle of explaining
something new, while accepting that there is nothing entirely new, and that
the roots of novelty in evolution lie in existing forms? As Dawkins shows in
The Ancestor’s Tale [2], humanity can be tracked back seamlessly to the first
replicating cells. Origins disappear in continuity.
The challenge of studying evolutionary change, for any lineage or character-
istic, is to steer a course between the Scylla and Charybdis of monotonous
continuity and elusive origin points. This holds true for human evolution as

& 2016 The Author(s) Published by the Royal Society. All rights reserved.
 Downloaded from [Link] on September 27, 2016

much as any other part of biodiversity. On the one hand, showing that hominins were making stone tools at 3.3 Ma, 2
there is continuity in many aspects of hominins back to the before the appearance of Homo as currently understood. In

[Link]
last common ancestor with apes, and on the other hand, this context it is worth remembering that, when Leakey
there are many novel traits that appear successively across et al. [6] described Homo habilis, they argued for abandoning
the subsequent five or more million years. The solution to the cerebral Rubicon so that the genus Homo could include
this difficulty that is explored in this themed issue is to a smaller brained stone tool maker. None of this undermines
focus on transitions. Evolution is about the change from the adaptive significance of changes in life history or body
one state (at whatever biological level) to another, which size, or tool-making, but shows that clear watershed points
demands a focus on the comparison of states across time, do not occur. Kimbel et al. [7] look back at the history of
or across organisms and their adaptations. Such transitions the Australopithecus-Homo divide and also come to the con-
can be major or minor, can be multiple or single, and can clusion that its significance has been exaggerated, and that
be related to the appearance and disappearance of whole it is better to think of small transitions accumulated over a

Phil. Trans. R. Soc. B 371: 20150229

taxa, or of particular traits. Human evolution is the sum of longer period (figure 1).
those transitions, and the papers here present new evidence One possibility is that the line is being drawn in the
and review some of these across the 5 million years of our wrong place, and that, as has been argued before, the earliest
lineage’s history. Although diverse and broad-ranging, 16 members of Homo are indeed adaptively closer to the austra-
papers cannot do justice to the whole range of issues involved lopithecines, and that early H. erectus represents the major
in the multiple transformations that have led from an ape-like transition [8]. Or even closer to the present, that the divide
ancestor to the modern world, but they do bring to the fore lies between all archaic hominins and modern humans.
some central and common themes. Certainly there are grounds for seeing H. erectus (or in some
taxonomic schemes H. ergaster; figure 2) as a grade shift rela-
tive to all earlier hominins, with biology and behaviour more
similar to modern humans than to earlier hominins.
2. Hominin evolution: Is there a major divide? A case can be made for each of these as a divide within
Football, it is often said, is a game of two halves. It is the hominin evolution, but a detailed examination of the evi-
same with human evolution, albeit of rather unequal lengths. dence suggests a much more diverse and cumulative
One half comprises the evolution of what are usually referred process. Antón et al. [9] show that H. erectus is highly vari-
to as the early hominins, those taxa that are closer to humans able, not just in body size (see [3]), but also in other aspects
than to living apes, but are generally placed in other genera of its morphology. The appearance of modern humans is
than Homo (Sahelanthropus, Ardipithecus, Ororrin, Kenyanthro- seen at about 195 Ka at Omo Kibish in Ethiopia (figure 3),
pus, Australopithecus and Paranthropus). These are highly but Stringer [10] argues that the lineage leading to it shows
diverse creatures, but are linked by showing a variety of derived traits earlier than this, and so the transition is
hominin-like traits (mostly related to dentition and inferred spread over several hundred thousand years, and that it is
locomotion), and an absence of the major markers of Homo, likely archaic populations persisted across the African
an enlarged brain and more complex behaviours. This half continent, some until into the late Pleistocene [11].
of human evolution is often referred to as the ‘bipedal Looking at the totality of hominin evolution, there is no
apes’, and occurs between approximately 7.0 and 2.8 Ma, broad division between the earlier and later phases, nor
when the first Homo appears in the record (or, depending between archaic and modern humans. These transitions are
on one’s view, approximately 1.5 Ma, when the last ‘bipedal significant, but the richer fossil record now in existence,
apes’ disappear). The second half, from between approxi- and the multiple techniques available for studying it, show
mately 2.8 Ma to the present day, is concerned with our that the major transitions of human evolution are comprised
own genus, and includes the evolution of modern humans of multiple smaller ones.
themselves. Here, the focus is on encephalization, changes
in life-history strategy, developing technology, expansion of
diet and geographical range and the emergence of cultural
processes of evolution. 3. Is simple gradualism the best model?
One of the relatively rare elements of this volume is that If there are no big or single step transitions in human evol-
both halves are included, so that rather than being seen as ution, does this mean that the evolutionary history of our
two different events, overlapping patterns can be observed. lineage is a straightforward case of unilineal, gradual
Jungers et al. [3], for example, consider body size across all change over millions of years? Is it a case of simple anagenetic
hominins and show that the early bipedal ape versus the change, with better adaptations replacing existing ones over
human-like hominins divide does not exist; they show over- time?
lapping patterns, and certainly no contrast between the two The evidence discussed in these papers cannot be said to
at a point of ‘origins’. The idea that the emergence of Homo support what was classically thought of as the punctuated
is associated with an increase in body size is not supported. equilibrium model of evolution, as proposed by Gould,
Other aspects of hominin biology also do not show a clear Eldredge, Vrba and others [12,13]. This model predicted
distinction between Homo and earlier hominins; Dean [4] that human evolution should consist of prolonged periods
shows that all hominins share some form of shift in life- of stasis, interspersed by sharp and rapid bursts of speciation
history relative to living apes, and the Homo-Australopithecus and adaptive change. That is clearly not the case, but does it
boundary exists but is not one that stands out strongly follow that what we see is simple gradualism?
among a range of life-history changes. It is not just a matter Almost certainly not. Spoor et al. [14] looked at the
of body size and maturation; Lewis and Harmand [5] discuss fossil evidence in East Africa between about 3.5 and 2.5 Ma,
their recent discoveries at Lomekwi in West Turkana, focusing on Kenyanthropus platyops. Their morphometric
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[Link]
Phil. Trans. R. Soc. B 371: 20150229
Figure 2. Homo erectus. Early representative (KNM-ER3733) of Homo erectus
(sometimes referred to as Homo ergaster), from Koobi Fora, East Turkana,
Kenya, discovered by Richard Leakey and his team in 1976. Photo credit:
Turkana Basin Institute/Richard Leakey. (Online version in colour.)
Figure 1. Early Homo. KNM-ER1470, from Koobi Fora, East Turkana. Discov-
ered by Richard Leakey and his team in 1967. Photo credit: Turkana Basin
high rates of transition. This suggests that hominin evolu-
Institute/Richard Leakey. (Online version in colour.)
tion is neither a simple punctuated process, nor a constant
gradual one, but a complex interaction between variable
study shows that K. platyops was different from both rates of change, environmental dynamics and the competi-
Australopithecus deyiremeda and A. afarensis, suggesting that tive interactions of the hominins and their sympatric
there were three contemporary taxa in east Africa during fellow-travellers in evolution.
the middle Pliocene. This mirrors the high level of diversity
that is known from the succeeding Plio-Pleistocene [15].
Even within the Pleistocene, as Antón [9] discusses for
H. erectus, and Stringer [10] and Mirazón Lahr [11] for H. 4. Is there a pattern to transitions? Geography
sapiens and its contemporaries, there is considerable diversity,
and clearly not a unilinear pattern. and the role of Africa
This suggests that human evolution is not just one taxon If the overall evidence suggests that hominin evolution
evolving from another, but involves branching and specia- includes multiple transitions, the next question to be posed
tion. This is not, of course, surprising, as it is how is whether there is any pattern to them. Perhaps the most
evolutionary change operates in other lineages, but it striking of these is the centrality of Africa in evolutionary
reinforces the similarity between patterns of human evolution change—the first hominins, the earliest diversification of
and those seen in other groups of animals. That this may be early hominins [14], the earliest stone tools [5], the earliest
part of the same process, subject to the same environmental Homo [7], the earliest Acheulean [17], the first modern
pressures as other animals, is hinted at by Fortelius et al. humans [10,11] and the beginnings of modern human behav-
[16] who analyse the Turkana Basin mammalian record, iour and organization [18,19]. There might be a taphonomic
one of the richest repositories of hominin fossils, as a ‘species bias in some of the dominance of Africa, but genetics,
factory’ between 4 and 2 Ma—particular combinations of which is not subject to such a bias, certainly supports it for
environmental conditions simultaneously creating refugia some of these events. While hominins were African, there is
and novel selective pressures ahead of their occurrence over a recurrent pattern of an origin in East Africa [14]; when
a wider area. hominins became global, the recurrent pattern is of an
The branching pattern of hominin evolution, the diversity African origin and dispersal beyond [9 –11]. Thus the adap-
at most periods, the lack of linear change and the survival of tive and contingent contexts for the transitions and early
archaic forms after their more derived descendants have phases of diversification of several lineages are to be found
evolved, all indicate something that is not simple phyletic in Africa, and perhaps East Africa more specifically. There
gradualism, to use Gould and Eldredge’s term [12]. Foley may well be stochastic elements involved in setting the
[15] looked at the overall patterns of hominin diversity over right initial conditions (the right sort of ape in the right sort
time and the appearance and disappearance of taxa (as of environment in Africa), but the repeated pattern merits
proxies for speciation and extinction), and found that while attention. General biogeographical properties must certainly
such events are distributed broadly across the whole of be involved (the tropics as areas of higher rates of speciation,
the sequence, there are some phases—in the Pliocene, the for example, and the interaction between glacial cycles and
Plio-Pleistocene and the later Pleistocene—when there were the relationship between Eurasia and Africa). The dynamic
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extinction of existing ones; they can be changes in mor- 4

phology, known mostly from the skeleton, especially the

[Link]
cranium and dentition; they can be changes in technology
and other aspects of behaviour inferred from the archaeologi-
cal record; and, increasingly, they can be changes in gene
distributions and frequencies, and the inferred lineages and
populations of ancient humans. How do these relate to
each other? Does behaviour (as attested in technology, for
example) precede morphological change? Or vice versa?
There is no simple answer to this question. On the one
hand, the appearance of stone tools by approximately
3.3 Ma [5] predates the known appearance of Homo by

Phil. Trans. R. Soc. B 371: 20150229

about half a million years and so also predates the expected
relationship between expanding brains and technology. On
the other hand, the appearance of the Acheulean appears to
be close to the first known H. erectus/ergaster [17], and so is
suggestive of a relationship. But in both cases there remains
room for doubt. The age of the earliest Homo was recently
Figure 3. The earliest representative of Homo sapiens from Omo Kibish,
increased from 2.3 to 2.78 Ma [22], and Kimbel [7] outlines
Ethiopia. Discovered by Richard Leakey and his team in 1967. Reproduced
the difficulties of pinpointing significant change; the age of
with permission from C. Stringer and M. Day. (Online version in colour.)
the first tools changed by nearly a million years with the dis-
coveries at Lomekwi. The evidence we currently have is
nature of the African environment over the period between much richer than even at the beginning of the twenty-first
3.5 and 2.0 Ma is discussed by Rose et al. [20], who show century, but it has still not stabilized to the point where
that the critical area in Africa, and its links to Eurasia (the such associations (or lack of them) can be regarded uncriti-
north-eastern quadrant), generates a distinctive biogeogra- cally; indeed, the Signor-Lipps effect [23] would suggest
phical pattern derived from the interaction between that it never could be.
orbitally driven factors (such as eccentricity and precession) The more recent parts of human evolution can offer
and local environmental conditions. Palaeoenvironmental insights. The broadly accepted time scale for the origins of
research in recent years has built greatly on the marine iso- the modern human lineage is thought to be between 0.4
tope revolution of the second half of the twentieth century, and 0.7 Ma, but there are grounds for recognizing transitions
by beginning to reveal the importance of local variation around and after this date. The dating is such that we can
within broader patterns and trends. This goes some way to resolve a number of important events within that time
explaining the variation between Turkana and other lake scale—a genetic divergence from the last common ancestors
basins in Africa, and even between East and West Turkana, with Neanderthals about 0.5 + 0.2 Ka, the appearance of
and how this translates into evolutionary dynamics [16]. prepared core technologies at approximately 0.3 Ka, the
These palaeoclimatic and palaeoecological studies also appearance of a modern morphology by at least 195 Ka, dis-
allow us to explore the relationships between the environ- persals across Africa at approximately 120 Ka and beyond
mental context and evolutionary change. Uno et al. [21] between 80–60 Ka or before, elements of modern human
highlight evidence from Turkana showing marked changes behaviour by at least 160 Ka, and various accelerations in
around 1.9 Ma, which, as they note, more or less coincides the accumulation of such traits by 120 Ka, and especially
with the appearance of the Acheulean, as described by de la after 100 Ka [10,11,18,19].
Torre [17]. Fortelius [16] also notes marked changes between Mirazón Lahr’s [11] detailed breakdown of the phases by
1.87 and 1.5 Ma, and this similarly coincides with a period which a small ancestral population in Africa was transformed
of evolutionary dynamism. These early phases suffer from a into a global, high density one, shows that when it is possible
lack of precise chronological resolution, so that the links to observe the evidence in fine detail, there was a complex
between specific events or transitions in hominin evolution micro-evolutionary interaction of population collapses and
and climate change cannot be made absolute, but this is expansions that continuously reshuffled diversity along a
more feasible for the last quarter of a million years, where lineage, and transformed the human species.
Mirazón Lahr [11] is able to link events in the evolution of
human diversity to a background of climate change.
There is no doubt that areas beyond Africa are also impor- 6. Is there a pattern to transitions? The causes
tant in human evolution—the dispersals of hominins into
Asia, the evolution of Neanderthals and Denisovans, the of transformations in human evolution
evolution of H. floresiensis—but the significance of Africa Placing the transitions towards becoming human into a
remains. chronological pattern and determining their environmental
contexts are the first steps in a broader scientific enterprise,
namely to gain insights into what caused these transform-
5. Is there a pattern to transitions? From ations. ‘Cause’ is, of course, a strong word, and by and
large studies of human evolution have not been short of
genotype to extended phenotype theories and models that attempt to find a single cause for
Changes in the course of hominin evolution take many the whole of it—an aquatic phase, seed-eating, hunting,
forms—they can be the evolution of new species, and the pair-bonding, hair-losing, tool-making and so on. As has
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already been discussed, there cannot be a single cause of their list, partly because humans possess a new means of 5
human evolution, because it consists of many independent information transmission (language and culture more gener-

[Link]
transitions. Indeed, any of the transitions is likely to involve ally), and thus potentially change the rules of evolution, and
multiple events that have many factors contributing to partly because we are so distinct compared with other
them, ranging from the changing climate, the immediate animals. Vinicius [26] added concepts of changing patterns
environment, local resource availability, and demographic, of modularity in the generation of complexity, leading to
cultural and social context. Biological factors, ranging from new information systems and higher levels of biological
genes to life-history strategies, will play a part, but so too organization. Foley [15] considers the question of whether
will behaviour. Bringing these all together is a major humans represent a major transition or not, focusing on the
challenge to the field. cumulative and dispersed nature of the events, seeing a pat-
Several of the papers address this, particularly for the tern that is consistent with many of the individual papers in
later part of the evolutionary time scale, and emphasize this volume. In terms of impact, and uniqueness among

Phil. Trans. R. Soc. B 371: 20150229

something that is often invisible for earlier periods—that extant animals, humans in the course of their evolution
demography and social group size and organization are cen- have undergone major changes, but the relatively good visi-
tral to human evolution. Marean [18] makes the points that it bility of the evolutionary record for humans (compared
is strong group structures and equally strong inter-group riv- to other manor transitions) shows that it consists of many
alry that are important derived human characteristics, and so smaller cumulative changes, not a single major transition.
the critical question is under what ecological conditions might
these occur in a way that transforms archaic human behav-
iour. Marean’s answer is that it is the rich and predictable
aquatic resources of southern Africa that—uniquely—provide 8. Future directions
these conditions. Tryon and Faith [19] pursue a similar argu- A volume such as this is a snapshot of current research about
ment in their paper in relation to the origins of the Later Stone human evolution, and strongly reflects recent discoveries and
Age in East Africa and propose that human adaptations may ideas. We hope that it will also serve a purpose in signposting
have been a response to coping with increased population future directions for research.
size or density. Key to both arguments is that it is the inter-
action between ecological conditions and the way in which (i) Multi-proxy approaches. Traditionally, research into
human social groups can distribute themselves across the human evolution has been compartmentalized, by
landscape that are at the heart of major changes, so providing approach, by time and by continent. Day-to-day this
a direct link between evolutionary transitions and the ecologi- may be inevitable, but the growth of multi-disciplin-
cal conditions in which hominins would have lived on the ary projects is encouraging. Theoretically, models of
landscape. Collard [24] extends this through the analysis of cultural evolution are becoming more and more impor-
ethnographic data to suggest that what is most likely to influ- tant, thus bringing archaeology into a more
ence human hunter–gatherer adaptation is environmental evolutionary framework; practically, material from
risk, and that many of the adaptations that develop in the sites can be subject to analyses from many different
course of human evolution, particularly technological technologies, bringing light to bear on human evol-
complexity, may be selected to reduce risk. ution as a whole. The answers to many of the
These examples focus on the foraging and ecological questions raised in this volume will come from greater
aspects of human adaptations, but these are just part of our integration across the disciplines involved, and wel-
overall phenotypic set, the traits that have emerged as critical coming further disciplines and technologies into the
to survival during the course of human evolution. Dean’s [4] field.
presentation of maturation patterns shows that shifts in life- (ii) New technologies and analytical methods. One of the uni-
history are likely to be equally important, and underlie fying threads across the papers in this volume is the
many of the behavioural and cultural adaptations, while enormous impact of scientific techniques—for dating,
Antón [9] reminds us that the gap between genetics and cul- for environmental reconstruction, for analysing mor-
ture is filled by many other possibilities, of which phology, for modelling behaviour. New ventures
phenotypic plasticity in response to variable environments is may start with finding a fossil or site, but it is the
an important one. application of new technologies that is transforming
We are a long way from understanding the nature of the the field. Beyond the sense of awe that we must feel
various transitions in human evolution, but we have made knowing the colour of the eyes of people at the end
considerable progress in recognizing that there is no single of the Upper Palaeolithic, there is the recognition
cause, as there is no single event, and there is no single that palaeogenomics can test many old ideas more
cause because many levels of biology and behaviour come thoroughly than ever before, and reach aspects of
into play. the past that were not accessible from observations
on fossils and stone tools alone.
(iii) The limits of concepts. We may like to think that
research answers questions and solidifies knowledge,
7. Is there a major transition? but it also opens up areas of uncertainty. This is true
Maynard Smith and Szathmary [25] proposed that across the across the whole range of palaeoanthropology—for
whole of evolution there have been a small number of major example, both the early hominins and the close rela-
transitions, ones where the rules of the biological world are tives of modern humans are exposing the difficulties
changed (for example, when sexual reproduction evolved), of thinking in terms of species as entities, thus pushing
and greater complexity emerged. Human evolution was on us towards more fluid models of how diversity
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evolves. The same holds true of archaeological con- 6

cepts, as entities such as the Oldowan come under

[Link]
scrutiny through the discovery of the even more
ancient lithics at Lomekwi. Future research is not
just a question of empirically filling in the gaps
around transitions, but also developing new concepts
and models to tackle a more resolved set of data.
(iv) The nature of hominin variation. In the relatively simple
world of palaeoanthropology of 50 years ago, vari-
ation in morphology reflected biology and genetics,
and variation in technology reflected culture. We are
now aware that there is much greater overlap and

Phil. Trans. R. Soc. B 371: 20150229

complexity. Stone tools may effectively be under
some level of genetic control (see Collard [27]), and
as Antón et al. discussed [9], morphological variation Figure 4. Richard Leakey in the field, with Kamoya Kimeu. Photo credit:
can reflect various levels of plasticity. But this in Turkana Basin Institute/Richard Leakey. (Online version in colour.)
turn raises questions about the genetic basis for plastic
responses and reaction norms. Future research will
have to grapple with a much closer and more complex ranging from the early Miocene to the early Pleistocene.
interaction between the biological and the behaviour- One of the key things about this palaeoanthropological pro-
al, the cultural and the genetic. The exciting prospect gress was the way in which fieldwork was interwoven with
is that new technologies may provide the tools to do building up institutions—first with the National Museums
so. of Kenya, then the Louis Leakey Memorial Institute, and
most recently the Turkana Basin Institute, with its field
stations on each side of the lake. Leakey was the first to recog-
9. Expanding the record nize the importance of providing an infrastructure in which
The papers in this volume are testimony to the growing not only his work, but the work of others, could thrive,
strength of new scientific methods, the breadth and inte- and, for a newly independent nation such as Kenya, capacity
gration of approaches, and the ability to place human building could begin. Such initiatives take vision, but they
evolution into the context of broader evolutionary concepts. also take practical organization and fund-raising, and over
These approaches have demonstrated remarkable power and the decades Leakey has brought tens of millions of dollars
offer equal potential. However, in the end, what we know into the field. Part of the skill of getting things done is to
about human evolution depends upon its fundamental basis, work well with others, and this has been a signature of
the fossil and archaeological records. It might seem that mod- Richard’s approach to the many paleoanthropologists and
elling, ancient genomes and three-dimensional morphometrics related scientists with whom he has worked—most notably
are the keys to the future, but all these depend on the continu- Glynn Isaac, Alan Walker and Bernard Wood. However, it
ing accumulation of new fossils and the discovery of new is not just a matter of Richard leading and others following,
archaeological and palaeontological sites. for there is also the crucial partnership with Meave Leakey,
Expanding the record is hard work, time consuming, fre- who continues the pattern of major discoveries that she and
quently arduous and is often considered to be high-risk by Richard began in the 1960s.
research funding bodies. Better to keep re-analysing existing This volume is devoted to exploring the major transitions
material than spend money on unknown areas of unknown in hominin evolution, and shows that it is a complex and
potential. There is often a long fallow period before signifi- cumulative process. As we developed the programme and
cant results come in—the case of Louis and Mary Leakey decided to focus on the major transitions we realized that
working for three decades at Olduvai Gorge before finding all of these were topics to which Richard Leakey and his col-
‘Zinjanthropus’ is a classic case in point. However, without leagues have contributed critical new finds. That we now
that personal and financial investment, there can be no pro- recognize this complexity is in no small part due to the dis-
gress in the field. Since the 1960s, that progress has been coveries that Richard and Meave Leakey have made. In any
remarkable, particularly in Africa. In 1958, there were five CV, it is not too bad to be able to list the revelation of the
or six accepted hominin taxa; now there are probably more diversity of early Miocene African apes [28,29], the earliest
than 25 recognized species, and seven genera have been known australopithecine [30], the diversity of early Homo
named. Many people have contributed to this expansion, [31], the earliest known African H. erectus [32], the most com-
but none more so than Richard Leakey, both directly and plete ancient hominin skeleton [33], the most complete early
through the support and assistance he has given to others paranthropine or robust australopithecine [34] and the first
(figure 4). Richard Leakey’s career and achievements go known anatomically modern human [35].
well beyond palaeoanthropology, but within the field they The Discussion Meeting sponsored by the Royal Society
are unparalleled. In the 1960s, he was instrumental in estab- and the British Academy on which these papers are based
lishing the Omo as a major palaeoanthropological field site, was in part an opportunity to celebrate Richard Leakey’s
swiftly followed by setting up the East Rudolf and Koobi major contributions to the field as well as tackle some of
Fora Projects. This rapidly yielded hominin fossils that revo- the ‘big’ issues in human evolution, and it is a great pleasure
lutionized the understanding of the early African radiations. and honour to dedicate this volume to him. We hope that his
In the 1980s, Richard Leakey extended work to the western achievements will continue to inspire new generations to go
side of Lake Turkana and produced spectacular fossils out and explore our past in all corners of the world.
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Competing interests. We declare we have no competing interests. the Discussion Meeting with great skill. This paper was improved 7
Funding. We received no funding for this study. by two anonymous reviewers. We would like to thank Jenna Lane
and the staff at the Royal Society for their help and very efficient

[Link]
Acknowledgements. We would like to thank the Royal Society and the
organization, and Helen Eaton for all the guidance she has provided
British Academy for their generous support. Clive Gamble, Hélène
in putting this issue together.
Roche, Mark Thomas and Bernard Wood chaired the sessions at

Editor biographies
Robert Foley is the Leverhulme Professor of Human Evolution at the University of Cambridge,
a Fellow of King’s College and a Fellow of the British Academy. He is a co-founder, with Marta
Mirazón Lahr, of the Leverhulme Centre of Human Evolutionary Studies at Cambridge, an inter-

Phil. Trans. R. Soc. B 371: 20150229

disciplinary research centre. His research has focused on the ecological basis of human evolution,
and in particular the application of Darwinian models to patterns and processes of human evol-
ution, ranging from the early hominins to recent human populations. He has worked extensively
in East Africa.

Lawrence Martin is Director of the Turkana Basin Institute and Professor of Anthropology at the
University of New York at Stony Brook. He is a palaeoprimatologist whose research focuses on
the evolution of hominoid primates and the development, structure and thickness of dental
enamel. He has conducted paleontological fieldwork at Miocene sites in Cameroon, Kenya, Pakistan
and Turkey. His laboratory research on enamel structure involves the use of confocal microscopy,
polarised light microscopy, and scanning electron microscopy. His research has addressed phylo-
geny reconstruction for extant and extinct hominoid primates, species recognition in the fossil
record and enamel development and thickness.

Marta Mirazón Lahr, a Fellow of Clare College, is a Reader in Human Evolutionary Biology and Direc-
tor of the Duckworth Laboratory at the University of Cambridge, where she and Robert Foley founded
the Leverhulme Centre for Human Evolutionary Studies. The focus of her research is the evolution and
diversity of our species, Homo sapiens. Her work involves a range of disciplines - human palaeontol-
ogy, evolutionary genetics, behavioural ecology, linguistics and prehistoric archaeology. She has
carried out fieldwork in the Amazon, the South Pacific, India, Oman, Libya and Kenya. She is the
director of the ERC-funded IN-AFRICA Project in Kenya, and a co-investigator in the Trans-Sahara
Project, and the African Genomes Project with the GeoGenetics Centre of Copenhagen.

Chris Stringer has worked at The Natural History Museum London since 1973 and is now Research
Leader in Human Origins and a Fellow of the Royal Society. His early research was on the relation-
ship of Neanderthals and early modern humans in Europe, but through his work on the ‘Recent
African Origin’ theory of modern human origins, he now collaborates with archaeologists, dating
specialists and geneticists in attempting to reconstruct the evolution of modern humans globally.
He has excavated at sites in Britain and abroad, and he is currently co-directing the Pathways to
Ancient Britain project, funded by the Calleva Foundation.
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