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RSTB_365_1556_Cover.qxd 9/7/10 10:42 AM Page 1

Phil. Trans. R. Soc. B | vol. 365 no. 1556 pp. 3263–3410 | 27 Oct 2010
ISSN 0962-8436

volume 365

27 October 2010 number 1556

volume 365 . number 1556 . pages 3263–3410
pages 3263–3410
The first four million years of human
evolution In this Issue

Papers of a Discussion Meeting issue organized and edited by Alan Walker and Chris Stringer The first four million years of human evolution
Introduction Papers of a Discussion Meeting issue organized and edited by Alan Walker and Chris Stringer
The first four million years of human evolution 3265
A. Walker & C. Stringer
Articles
In search of the last common ancestor: new findings on wild chimpanzees 3267
W. C. McGrew
More reliable estimates of divergence times in Pan using complete mtDNA sequences
and accounting for population structure 3277
A. C. Stone, F. U. Battistuzzi, L. S. Kubatko, G. H. Perry Jr, E. Trudeau, H. Lin & S. Kumar
Spinopelvic pathways to bipedality: why no hominids ever relied on a bent-hip–bent-knee gait 3289
C. O. Lovejoy & M. A. McCollum
Arboreality, terrestriality and bipedalism 3301
R. H. Crompton, W. I. Sellers & S. K. S. Thorpe
Two new Mio-Pliocene Chadian hominids enlighten Charles Darwin’s 1871 prediction 3315
M. Brunet

The first four million years of human evolution

Phylogeny of early Australopithecus: new fossil evidence from the Woranso-Mille (central Afar, Ethiopia) 3323
Y. Haile-Selassie
Anterior dental evolution in the Australopithecus anamensis–afarensis lineage 3333
C. V. Ward, J. M. Plavcan & F. K. Manthi
Molar microwear textures and the diets of Australopithecus anamensis and Australopithecus afarensis 3345
P. S. Ungar, R. S. Scott, F. E. Grine & M. F. Teaford
An enlarged postcranial sample confirms Australopithecus afarensis dimorphism was similar
to modern humans 3355
P. L. Reno, M. A. McCollum, R. S. Meindl & C. O. Lovejoy
The cranial base of Australopithecus afarensis: new insights from the female skull 3365
W. H. Kimbel & Y. Rak
Hominin diversity in the Middle Pliocene of eastern Africa: the maxilla of KNM-WT 40000 3377
F. Spoor, M. G. Leakey & L. N. Leakey
Stable isotopes in fossil hominin tooth enamel suggest a fundamental dietary shift in the Pliocene 3389
J. A. Lee-Thorp, M. Sponheimer, B. H. Passey, D. J. de Ruiter & T. E. Cerling
Retrieving chronological age from dental remains of early fossil hominins to reconstruct human
growth in the past 3397
M. C. Dean

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The first four million years of human evolution
Papers of a Discussion Meeting held at the Royal Society on 19 and 20 October 2009.
Organized and edited by Alan Walker and Chris Stringer

Contents

Introduction
The first four million years of human evolution 3265
A. Walker and C. Stringer

Articles
In search of the last common ancestor: new findings on wild chimpanzees 3267
W. C. McGrew

More reliable estimates of divergence times in Pan using complete mtDNA sequences 3277
and accounting for population structure
A. C. Stone, F. U. Battistuzzi, L. S. Kubatko, G. H. Perry Jr, E. Trudeau, H. Lin
and S. Kumar

Spinopelvic pathways to bipedality: why no hominids ever relied on a 3289

bent-hip – bent-knee gait
C. O. Lovejoy and M. A. McCollum

Arboreality, terrestriality and bipedalism 3301

R. H. Crompton, W. I. Sellers and S. K. S. Thorpe

Two new Mio-Pliocene Chadian hominids enlighten Charles Darwin’s 1871 prediction 3315
M. Brunet

Phylogeny of early Australopithecus: new fossil evidence from the Woranso-Mille 3323
(central Afar, Ethiopia)
Y. Haile-Selassie

Anterior dental evolution in the Australopithecus anamensis – afarensis lineage 3333

C. V. Ward, J. M. Plavcan and F. K. Manthi

Molar microwear textures and the diets of Australopithecus anamensis and 3345
Australopithecus afarensis
P. S. Ungar, R. S. Scott, F. E. Grine and M. F. Teaford

An enlarged postcranial sample confirms Australopithecus afarensis dimorphism 3355

was similar to modern humans
P. L. Reno, M. A. McCollum, R. S. Meindl and C. O. Lovejoy

The cranial base of Australopithecus afarensis: new insights from the female skull 3365
W. H. Kimbel and Y. Rak

Hominin diversity in the Middle Pliocene of eastern Africa: the maxilla of 3377
KNM-WT 40000
F. Spoor, M. G. Leakey and L. N. Leakey

3263
3264 Contents

Stable isotopes in fossil hominin tooth enamel suggest a fundamental dietary 3389
shift in the Pliocene
J. A. Lee-Thorp, M. Sponheimer, B. H. Passey, D. J. de Ruiter and T. E. Cerling

Retrieving chronological age from dental remains of early fossil hominins to 3397
reconstruct human growth in the past
M. C. Dean
 Downloaded from rstb.royalsocietypublishing.org on October 24, 2010

Phil. Trans. R. Soc. B (2010) 365, 3265–3266

doi:10.1098/rstb.2010.0179

Introduction

The first four million years of human

evolution

In one of the last paragraphs of The origin of species As discussed in one of the first contributions to this
(1859), Darwin famously suggested that ‘Much light volume, molecular estimates of the divergence time
will be thrown on the origin of man and his history’. between humans and chimpanzees presently converge
When he published The descent of man 12 years on approximately 5– 7 Ma, although we two are old
later, there was still no fossil evidence of our earliest enough to remember the pre-molecular days, when
evolutionary history, and nothing at all from the the supposed uniqueness of humans seemed to require
African continent. Yet our close biological relationship a time-span 2– 3 times those figures to account for the
to the great apes, and especially the African apes, the evolution of special features like bipedalism and high
gorilla (Gorilla gorilla) and chimpanzees, had long encephalization. But now, fossils of putative human
been recognized, even by scientists who were ignorant lineage members have been reported from approxi-
of, or unsympathetic to, evolutionary thinking. mately 6 Ma deposits in Chad and Kenya, and fossils
Nevertheless, when we remember his cautious nature of the genus Ardipithecus from approximately 4.4 Ma
and the continuing powerful opposition to his ideas, sediments in Ethiopia include about 40 per cent of a
it still required fortitude for Darwin to venture ‘It is complete skeleton. Views differ on the relationship
therefore probable that Africa was formerly inhabited of these forms to each other, and to the succeeding
by extinct apes closely allied to the gorilla and chim- and better-known genus Australopithecus. Several
panzee; and as these two species are now man’s skeletons of the latter have been found in the last
nearest allies, it is somewhat more probable that our few years. These include an adult from Sterkfontein
early progenitors lived on the African continent than cave, South Africa, not yet certainly dated, another
elsewhere’. In explaining why the fossil evidence of adult from 3.8 Ma deposits in Woranso-Mille,
our origins was slow to appear, he prophetically Ethiopia, a 3.3 Ma child’s skeleton from Dikika,
stated ‘Nor should it be forgotten that those regions Ethiopia and four partial skeletons from Malapa
which are the most likely to afford remains connecting Cave, South Africa, dated to about 1.9 Ma. Dozens
man with some extinct ape-like creature, have not as of other less complete hominin fossils from approxi-
yet been searched by geologists’. In fact it was to mately 6 to 2 Ma have been found, as well as these
take another 50 years before such fossil evidence skeletons.
began to emerge in Africa itself, and Darwin would Our meeting was timed to coincide with the double
have been amazed by the remarkable evidence which celebration of Darwin’s 200th birthday and the 150th
has accumulated since then concerning the earliest anniversary of the publication of The Origin of Species,
stages of human evolution. and to take the first opportunity to bring together as
Spectacular discoveries of early members of the much as possible of the rich, newly published data
human lineage, including nearly complete skeletons concerning the earliest-known members of the
and dozens of other 6 to 2 Ma fossils have been human lineage. Through the generosity of the partici-
made in the last 10 – 20 years. Single complete skel- pants, our hope that detailed images and casts of the
etons are much more useful analytically than new material would be brought together for the first
separate parts of many individuals, yet until recently, time during the meeting was amply met, although in
few had been found from the period before 2 Ma. the event only one of us could be there to see the
Even Australopithecus, discovered in South Africa outcome.
in 1924, and published and named in 1925, is still The meeting was also planned to showcase the
relatively incompletely known. For instance, the interdisciplinary nature of palaeoanthropology, by
famous ‘Lucy’ skeleton from Ethiopia is only about highlighting the new methods that have been devel-
20 per cent intact. But new and more complete early oped to extract behavioural and life history
hominin skeletons from different parts of the African information from fossils. These included computer
continent now promise to give us a much more modelling of locomotor capabilities, finite element
complete picture of the early phases in the history of modelling of stresses in bone, laser scanning compari-
the human lineage. sons of joint surfaces, quantification of semicircular
canal morphology and its relationship to head
motion, isotope analysis of teeth for dietary and cli-
mate reconstruction, confocal microscopy and
texture analysis of tooth wear to indicate diet, and
One contribution of 14 to a Discussion Meeting Issue ‘The first four
million years of human evolution’. reconstruction of life history parameters from

3265 This journal is # 2010 The Royal Society

 Downloaded from rstb.royalsocietypublishing.org on October 24, 2010

3266 A. Walker & C. Stringer Introduction

incremental lines in tooth enamel and dentine. The new material such as the just-published Ardipithecus
analytical sessions highlighted what could be accom- skeleton, and the reconstructed Sahelanthropus
plished by the careful reconstruction, study and cranium.
analysis of the new fossils. In the event, we have not been able to publish all of
By concentrating on the early part of the record of the contributions made at the meeting, and this unfor-
human evolution, the meeting was also able to docu- tunately included a description of the very complete
ment the essential ecological, behavioural, and australopithecine skeleton from Sterkfontein men-
morphological stages that underpinned the subsequent tioned earlier. Nevertheless we feel that The first four
emergence of the genus Homo. Field workers reported million years of human evolution was an appropriate
on studies of the behaviour of wild chimpanzees as measure of how much progress the field of palaeoan-
possible models for early hominin behaviour, and on thropology (a term unknown 150 years ago) has
the geological and environmental setting of the fossils, made in meeting Charles Darwin’s expectations. We
as well as their anatomy and preservation. Context for would like to thank all the staff of the Royal Society
the discoveries was provided by colleagues who, for who worked on the planning and running of the meet-
example, used tephrostratigraphy, argon – argon radio- ing, and the editorial team who has worked so hard to
metric dating, faunal and floral analysis, GIS satellite bring this volume to fruition.
imagery and taphonomy.
Our hope was to bring about a new understand-
ing of early hominin evolution by bringing together Alan Walker1
the newest fossils and the latest analytical methods, Chris Stringer2,* June 2010
and we think the meeting at least helped progress
1
towards that ambitious goal. But the meeting also Anthropology & Biology, Penn State University,
provided the first opportunity to present many of University Park, PA 16802, USA
2
the newest discoveries to scientific and public audi- Dept of Palaeontology, The Natural History Museum,
ences alike. A memorable conference dinner was London SW7 5BD, UK
accompanied by a display of replicas of spectacular *Author for correspondence ([email protected]).

Phil. Trans. R. Soc. B (2010)

 Downloaded from rstb.royalsocietypublishing.org on October 24, 2010

Phil. Trans. R. Soc. B (2010) 365, 3267–3276

doi:10.1098/rstb.2010.0067

In search of the last common ancestor: new

findings on wild chimpanzees
W. C. McGrew*
Leverhulme Centre for Human Evolutionary Studies, Department of Biological Anthropology,
University of Cambridge, Cambridge CB2 1QH, UK
Modelling the behaviour of extinct hominins is essential in order to devise useful hypotheses of our
species’ evolutionary origins for testing in the palaeontological and archaeological records. One
approach is to model the last common ancestor (LCA) of living apes and humans, based on current
ethological and ecological knowledge of our closest living relations. Such referential modelling is
based on rigorous, ongoing field studies of the chimpanzee (Pan troglodytes) and the bonobo (Pan
paniscus). This paper reviews recent findings from nature, focusing on those with direct implications
for hominin evolution, e.g. apes, using elementary technology to access basic resources such as food
and water, or sheltering in caves or bathing as thermoregulatory adaptations. I give preference to
studies that directly address key issues, such as whether stone artefacts are detectible before the Old-
owan, based on the percussive technology of hammer and anvil use by living apes. Detailed
comparative studies of chimpanzees living in varied habitats, from rainforest to savannah, reveal
that some behavioural patterns are universal (e.g. shelter construction), while others show
marked (e.g. extractive foraging) or nuanced (e.g. courtship) cross-populational variation. These
findings allow us to distinguish between retained, primitive traits of the LCA versus derived ones
in the human lineage.
Keywords: tool use; shelter; diet; ranging; last common ancestor; chimpanzee

1. INTRODUCTION the extinct LCA, based on indirect evidence. Thus,

This paper aims to synthesize and to update recent this synthesis covers technology, diet, shelter and
(from 2005 onwards) findings from studies of the ranging and foraging.
ethology and ecology of wild chimpanzees (Pan Attempts to use findings from ethological and eco-
troglodytes) that are relevant to modelling human logical (as opposed to morphological) research on
origins. Given space constraints, this exercise will be chimpanzees to model the behaviour of ancestral
limited to field studies, and therefore mostly to obser- humans are relatively recent, dating from the rise of
vational data on the spontaneous behaviour of apes primatological field studies in the last 50 years.
in situ, cited selectively. It emphasizes primary reports, Although most early field workers were interested in
usually journal articles, on the assumption that older apes in their own right, their mentors often had in
secondary reviews (e.g. Mitani et al. 2002; McGrew mind the potential applicability of the exciting new
2004) provide access to earlier material. It concen- findings to human issues (e.g. Goodall & Hamburg
trates on the eight study sites with fully habituated 1974). Many of the early attempts now look crude
subjects, here listed in the order of seniority: Gombe and simplistic (e.g. McGrew 1981). For example,
(Tanzania), Budongo (Uganda), Mahale (Tanzania), most were content to talk about extinct hominids as
Kanyawara (Uganda), Bossou (Guinea), Taı̈ (Ivory a single unspecified class, but as the hominin evol-
Coast), Ngogo (Uganda) and Fongoli (Senegal). utionary record became more and more diverse, with
However, given the geographical bias to eastern and more and more taxa unearthed, this monolithic exer-
western Africa, other sites with partly habituated sub- cise was less and less satisfactory. Furthermore, as
jects, especially in central Africa, such as Goualougo data began to emerge on wild bonobos, Pan paniscus
(Republic of Congo), are necessarily invoked too. (Kano 1992), who are as equally closely related as
Most importantly, it focuses on topics that are relevant chimpanzees to hominins, and as cross-populational
to modelling the behaviour of the last common ances- variation began to emerge in chimpanzees (McGrew
tor (LCA) of the divergent lines that led to living 1992), easy generalizations grew harder to make. Eco-
humans and living chimpanzees. These topics are pre- logical studies of chimpanzees in a variety of ecotypes,
sented in terms of their ‘directness’ in comparisons from rainforest to savannah, forced more precise mod-
between what primatologists see now in living apes, elling (Moore 1996). Finally, debate over the best way
and what palaeoanthropologists seek to infer about to model human origins and evolution, that is, via
referential versus strategic models, or by homology
versus analogy, muddied the waters (e.g. Tooby &
*[email protected] DeVore 1987). Sayers & Lovejoy (2008) took the
One contribution of 14 to a Discussion Meeting Issue ‘The first four extreme position that chimpanzees may be no more
million years of human evolution’. useful as models than other, more ecologically,
3267 This journal is q 2010 The Royal Society
 Downloaded from rstb.royalsocietypublishing.org on October 24, 2010

3268 W. C. McGrew Chimpanzees and the last common ancestor

zoogeographically and phylogenetically distant taxa, innovation in wild chimpanzees, at Mahale, shows
such as capuchin monkeys (Cebus spp.). More recently, inventiveness to be common, but the chance that a
Lovejoy (2009) has asserted that extant African ape- novel behavioural pattern will be propagated and
based models are no longer appropriate (for a contrary become established in a population is rare (Nishida
view, see Whiten et al. 2010). However, for focused et al. 2009).
studies, such as of Oldowan lithic industries, apes No longer is it enough just to list the types of tool
may still be the model of choice (Toth & Schick 2009). found at a given site, as nominal (presence/absence)
This paper takes the conservative line that lacking data. Now attention to relative frequency and compe-
a fossil record for apes since the Miocene (cf. tence of performance across age and sex classes is
McBrearty & Jablonski 2005), and having only a expected, along with data on context, variation in
shallow archaeological record for apes, all that we form and function of the tools’ manufacture and use
sensibly can hope to model is the LCA. In doing so, (e.g. Sanz et al. 2009a). Functional (e.g. extractive
I make several simplifying assumptions, such as that foraging), biomechanical (e.g. percussive) and cogni-
anything that a chimpanzee can do today, the LCA tive (e.g. artefact complexity) aspects of technology
could have done 6 – 7 Myr ago. Another pragmatic are stressed. Anecdotal versus idiosyncratic versus
assumption is that although the LCA could have habitual use of tools is differentiated. Distinction is
resembled the living chimpanzee, or bonobo, or drawn between a tool kit (i.e. the whole repertoire of
neither, or some combination of the two, most of a community’s collective range of tools) and a tool
what we have to work with on grounds of homology set (i.e. the obligate sequence of two or more tools
comes from P. troglodytes. Therefore, until comparable used to achieve a single goal). Composite tools (i.e.
breadth and depth of data are available for P. paniscus, when two or more objects are used simultaneously
the chimpanzee must carry the load. and complementarily to achieve a goal), such as
hammer and anvil (Carvalho et al. 2009), are distin-
guished from compound tools (i.e. when two or
(a) Technology more elements of different types are combined into a
Most of what behavioural primatologists have to offer single unit), such as a wedge used to level an anvil’s
to palaeoanthropology relies on artefacts, as these working surface (Biro et al. in press). Typology is
objects are comparable to what is found in the archae- now part of chimpanzee technology.
ological record. However, artefacts are the products of Tool kits show both uniformity and variety across
behaviour, and sometimes archaeological data are a populations. Sanz & Morgan (2007) presented quanti-
further step removed: butchery cutmarks on bones tative and qualitative findings from the Goualougo,
are the products of the ephemeral acts that produced Republic of Congo, chimpanzees, whose tool kit num-
them. Whatever the caveats, primatologists can offer bers 22 types, of which nine are used habitually
something that no archaeologist will ever see, that is, (customary). In contrast, Watts (2008b) published
BOTH the product AND the behaviour, directly comparable data from Ngogo, Uganda, where the
recorded. When a glancing blow of a stone hammer total tool kit numbers only 10 types, with four of
being used to crack a nut hits instead the stone anvil, these being habitual. Such variation suggests the possi-
producing a conchoidally fractured flake, the observer bility of a normally distributed spectrum, but this is
can see whether this was an accident. An archaeologist not the case. As with Goualougo, all habituated popu-
given only that same single flake could draw no valid lations show about the same-sized tool kits: Gombe
inference about the percussionist’s intentions. (22), Bossou (21), Taı̈ (21) and Mahale (16). How-
Studies of tool use by apes in nature have come ever, along with Ngogo, the other Ugandan sites
a long way from piecemeal natural history notes show small tool kits: Budongo (8) and Kanyawara
collected opportunistically and descriptively, to com- (10) (Sanz & Morgan 2007, table 3). Even more strik-
prehensive, systematic, hypothesis-driven empirical ing is the contrast between Goualougo and Ngogo
efforts, some of which are experimental. Comparative with regard to the predominate types of tools: the
analyses of chimpanzee material culture are done at top three at Goualougo are used in subsistence, that
every level, of individuals, lineages, communities, is, extractive foraging of termites, honey and water;
populations, subspecies and species (McGrew 2004). the top three at Ngogo are used in hygiene, especially
The chimpanzee ethnographic record now spans so wiping the penis after copulation, and in courtship.
many study sites across equatorial Africa that even (The reverse is equally true: Goualougo chimpanzees
chimpologists have trouble keeping them straight. very rarely use napkins, and Ngogo chimpanzees
Although only eight sites consistently allow all-day, rarely harvest insects.) However, some types of tool
close-up observation, there are five times as many use are chimpanzee universals, being found in all
other sites with varying degrees of habituation. In the long-studied populations across Africa, such as leaf
5 years, long-term sites studying the central (Hicks sponge (drinking water), aimed throw (weapon), play
et al. 2005; Sanz & Morgan 2007) and Nigerian start (toy), branch drag (display), etc.
(Fowler & Sommer 2007) subspecies have joined the Of particular importance is percussive technology,
longer term studies in eastern and western Africa. that is, the application of ballistic force via one
Even sites that have yet to habituate their subjects object to another to achieve a goal (Ling et al. 2009).
have yielded new behavioural patterns, e.g. root- In chimpanzees, this most famously takes the form of
digging at Ugalla, Tanzania (Hernandez-Aguilar hammer and anvil used to crack nuts, but it also
et al. 2007), fruit-cleaving at Nimba, Guinea (Koops occurs in smashing hard-shelled objects directly
et al. 2010), etc. The only comprehensive study of against anvils, in agonistic clubbing of adversaries or
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Chimpanzees and the last common ancestor W. C. McGrew 3269

in display, or in specialized extractive foraging such as of compound tool (Sousa et al. 2009), but it barely
pestle-pounding (Yamakoshi & Sugiyama 1995). In qualifies, being iterative. The most obvious example
the latter case, the pestle is a detached palm frond, of compound technology (albeit not tool use) in
the mortar is the apical growth tip of an oil palm non-human primates in nature is the sleeping plat-
(Elaeis guineensis) and the result is a cavity full of forms/nests/beds that are woven daily by great apes
mashed-up slurry, which is eaten. Anvil use without (see below). The best-known example in the extractive
hammers occurs when a hand-held, hard-shelled foraging of chimpanzees is the anvil– wedge, known
fruit is bashed directly against a boulder or root, as only from the nut-cracking of the Bossou chimpanzees
with baobabs (Adansonia digitata). Marchant & (Matsuzawa 2006). Bossou’s stone anvils are movable,
McGrew (2004) hypothesized an evolutionary scen- and so their positioning can be adjusted; anvils with
ario that led from anvil use to stone-knapping. near-horizontal working surfaces are the most efficient,
Tool sets in apes were first recognized in honey as the yielded nut-meat is readily picked up. An angu-
extraction (Brewer & McGrew 1990). In seeking to lar anvil can be levelled by inserting a smaller stone
harvest nature’s most calorific food, the minimal tool as a wedge underneath, to make the working surface
set requires a tool to break into the bees’ storage reser- less tilted.
voir and another tool to extract the liquid. That is, To what extent is the technological repertoire of the
some kind of percussive tool, such as hammer or chimpanzee now known? The steepness of the cumu-
chisel, plus some kind of dip-stick, are needed to lative ethnographic curve may be less than in the last
secure the food item (for the most complete treatment century, but it has not flattened out. New habitual pat-
of this resource’s exploitation, see Sanz & Morgan terns continue to be described: chimpanzees use spears
2009). Tool sets may be more complex: Boesch et al. to skewer small mammals (Pruetz & Bertolani 2007)
(2009) recently described tool sets used by the chim- and digging sticks to unearth roots (Hernandez-Aguilar
panzees of Loango, Gabon, in which up to five tools et al. 2007). Furthermore, new modes of tool use
were needed, e.g. pounder, perforator, enlarger, col- continue to emerge, such as the chimpanzees of
lector and swab. Tool sets also are used to exploit Nimba, Guinea, using cleavers to break apart large,
other animal prey, e.g. termites (Deblauwe et al. fibrous Treculia fruits (Koops et al. 2010).
2006; Sanz & Morgan 2007), ants (Sanz et al. Much progress has also been made on how individ-
2009b) and even when getting honey, the protein- ual apes in nature learn to use elementary technology.
aceous bonus of bee brood may be important too. Previous studies were descriptive or qualitative,
The key point about a tool set is that it is sequential whereas modern ones use sophisticated multivariate
task: if an A –B – C – D is necessary, then A – C– B – D analyses (e.g. general linear mixed model) to tease
will not do; you cannot check the oil level in your out the influences of independent variables.
car’s engine via the dip-stick, until you have opened Lonsdorf ’s (2006) study of termite fishing at Gombe
the car’s bonnet. Although tool sets may suggest showed that although all chimpanzees in the Kasakela
advanced cognitive abilities, many such mandatory community show this tool use by 5.5 years of age,
sequences are shown by creatures with modest brains daughters acquire the skills earlier, and this acquisition
(Hansell 2004), especially in shelter construction is a function of the mother’s overall time spent in the
(see below). What is impressive (and possibly activity. Humle et al. (2009) showed that chimpanzee
unique) about chimpanzee tool sets is that alternative infants at Bossou who had more opportunities to
versions may be used flexibly by different apes to observe their mothers started ant-dipping sooner and
solve the same problem. were more proficient than their low-opportunity
In human elementary technology, composite tools counterparts. However, in neither case were individual
are well known: Mortar and pestle, bow and arrow, differences in mother’s performance reflected in individ-
etc. Each element may stand alone, but is almost use- ual differences in their offspring, nor was there any
less without its partner. (Tool composites differ from direct teaching by mothers. Youngsters learned to fish
tool sets in that they are used simultaneously, rather or to dip by passive observational learning of tolerant
than sequentially.) Tool composites are known for models. Matsuzawa et al. (2001) have termed this
apes (see summary in Sugiyama 1997), and some are dyadic conduit of information from one ape to another
widespread, for example, in all populations where as ‘education by master–apprenticeship’.
chimpanzees use long wands to dip for driver ants, Some primatologists now apply archaeological
they also use bent-over saplings as a perch while methods to the study of chimpanzee technology in
doing so, to avoid the painful bites of the ants swarm- nature. Mercader et al. (2002, 2007) have shown that
ing on the ground below (McGrew 1974). However, the past nut-cracking activities of the Taı̈ chimpanzees
only recently have tool composites been systematically leave behind a record of stone artefacts. These can be
studied: Carvalho et al. (2009) showed that certain distinguished from human artefacts or naturally splin-
combinations of stone hammers and anvils were used tered rocks by ‘blind’ assessors, dated by standard
over and over again by the chimpanzees of Bossou, radiometric techniques (C14), and yield organic resi-
even taking into account the apes’ separate, indepen- dues (starch grains) that reveal their function. We
dent preferences for hammer or anvil. can now speak of a chimpanzee ‘stone age’ with time
Compound tools are harder to find in living apes in depth. Carvalho et al. (2008) applied one of the core
nature, although their production is readily induced concepts of archaeology, the chaine operatoire, to the
under contrived captive conditions. Combination of nut-cracking of Bossou’s chimpanzees, showing that
multiple items of the same type, e.g. leaves compressed from start to finish, this analytical technique is equally
together in leaf-sponging for water, is the simplest kind applicable to apes as to humans. Even retrospective
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3270 W. C. McGrew Chimpanzees and the last common ancestor

analyses of chimpanzee artefacts, in this case the but one of the keenest hunting populations, the chim-
brush-sticks used to fish for termites, as found in a panzees of Taı̈, does not hunt the small forest
museum, may explain how they were made (Heaton & antelopes (Cephalophus spp.) that are plentiful there.
Pickering 2006; cf. Sanz & Morgan 2007). The At the same time, several populations of bonobos
extent to which the archaeology of non-humans can avidly hunt antelope, but were said to show no interest
be pushed back in time remains to be seen, but a in primate prey; this apparent species difference evap-
new field is now underway (Haslam et al. 2009). orated with Surbeck & Hohmann’s (2008) report that
Finally, here is a sobering thought: of all the tools the bonobos of Lui Kotale, Democratic Republic of
named so far, only some of the hammers and anvils, Congo, also hunt guenons (Cercopithecus spp.). What
and some of the missiles thrown, will have a chance differs between the two sibling species of chimpanzee
of persisting in the archaeological record, taphonomy and bonobo is the sexual politics of meat-sharing.
willing, because they are made of stone. All of the In bonobos, females control the carcass and
others are made of organic raw materials, e.g. plant distribute the meat, and their collective dominance
or animal matter, and so will perish over time. There over males sometimes leaves the males with none,
are other lithic objects used, e.g. stones in self-tickle, even if individually a male can dominate a female
pebbles in play start, boulders in splash display, etc., (Hohmann & Fruth 2008). In chimpanzees, males
but it is unlikely that these will be archaeologically often control carcasses, and there has been much
recognizable. debate about how the sharing of the meat functions
in chimpanzee society. Now come solid data to test
Stanford’s (1999) hypothesis of meat-for-sex, that is,
(b) Diet that males selectively give meat to females in exchange
The chimpanzee is an omnivore, as all well-studied for sexual favours. Gomes & Boesch (2009) report that
populations show a mix of herbivory and faunivory. females copulate more often with males who share
The former is dominated by ripe fruit, but also meat with them in the long term. Thus, the female
includes leaves, pith, seeds, flowers, bark, gum, etc. need not be in oestrus at the time of the hunt, but
The latter focuses on social insects (ants, bees, rather forms a relationship that mutually enhances
termites) and small- to medium-sized mammals, the lifetime reproductive success of male (insemination
especially monkeys. Invertebrates usually are taken by probability) and female (nutritional enhancement).
tool-assisted extractive foraging, such as ant-dipping, However, meat-sharing in some chimpanzee
ant-fishing, honey-dipping and termite-fishing, that populations, e.g. Gombe in Tanzania (Gilby 2006),
is, by gathering. Until recently, vertebrate prey were appears to be driven by different mechanisms:
known to be captured and dispatched only by hand, intimidation, harassment, reciprocity, etc. Less likely
without technology. Pruetz & Bertolani (2007) is Tennie et al.’s (2009) ‘meat-scrap’ hypothesis that
showed that the chimpanzees of Fongoli, Senegal, use meat-sharing can be explained by the micro-nutrients
a weapon-assisted hunting technique to disable or kill found in even small amounts of meat. Meat-eating is
bushbabies while they sleep during the day in tree only one kind of faunivory, and the same nutrients
holes. The weapon is a sharpened stick (spear), can be easily obtained from invertebrates, which
jammed into the prosimian’s sleeping chamber. (Some chimpanzees eat daily.
sceptics have questioned whether the technique quali- Male sharing of prized foodstuffs with females also
fies as hunting, or the instrument as a spear. When an occurs with plant foods, which otherwise is rare in
Inuit waits beside a seal’s air-hole in the ice, then thrusts apes, usually occurring only between mother and
a sharp-ended linear object into it, skewering the prey, infant. However, Hockings et al. (2007) showed that
we are happy to call it hunting, so why not for apes?). when males at Bossou raided crops, especially
Notably absent from the diets of most chimpanzee papaya (Carica papaya), they almost always shared
populations are the underground storage organs the proceeds with females of reproductive age, even
(USO) of plants, that is, bulbs, roots, tubers, corms, when the latter were not in oestrus. These sharing pat-
rhizomes, etc. This absence was thought to reflect the terns reflected patterns of later sexual consortship.
generalized, non-digging hands of primates, plus the What about scavenging? Scattered, anecdotal reports
apes’ lack of the appropriate technology, that is, the dig- of chimpanzee scavenging mammalian prey have
ging stick. Hernandez-Aguilar et al. (2007) recently appeared from time to time, but no systematic study
described how the chimpanzees of Ugalla, Tanzania, was done until Watts (2008a) documented all known
dig up roots, using sticks and pieces of bark that show scavenging opportunities at Ngogo over 11 years of
the abraded wear patterns of repeatedly used digging observations totalling over 10 000 h. In that period, he
tools. Spat-out wadges of fibrous roots show them to saw only four scavenging events, and opportunities
be chewed and sucked, then discarded. A similar pro- were rare, occurring on average only every 100 h. This
cessing technique is used by the chimpanzees of contrasts mightily with over 650 kills made in over 270
Tongo, Democratic Republic of Congo, to get drinking hunts in the same period (Watts & Mitani 2002). Similar
water from subterranean tubers, but these are dug up pictures of rarity emerge from Gombe, Mahale and Taı̈.
by hand from friable, volcanic soils (Lanjouw 2002). Chimpanzees are not scavengers, it seems.
Across the continent, from Tanzania to Ivory Coast,
chimpanzee hunters take more monkeys as prey than
all other types of vertebrates combined, especially (c) Shelter
favouring the red colobus (Piliocolobus badius) (e.g. Shelter can be defined as the use of any material object
Watts & Mitani 2002). Others also hunt ungulates, to buffer the effects of the elements. A universal
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Chimpanzees and the last common ancestor W. C. McGrew 3271

behavioural pattern among great apes is their daily as the typical ecotype for wild chimpanzees, and so
construction of arboreal sleeping platforms: every contrast their ecological context with that of hominins
weaned individual builds an overnight nest and many who lived in more seasonal, mosaic habitats, this
also build day nests for napping. These compound restrictive picture is less and less tenable. Most of the
artefacts are scattered over the landscape and may study sites at which chimpanzees have been studied,
endure for months, leaving a record of points in and at least (depending on definition) three (Fongoli,
space where chimpanzees spend most of their lives. Gombe and Mahale) of the eight where the apes
(Chimpanzees typically retire at dusk and arise at have been fully habituated to close-range observation,
dawn, and so spend half of each tropical circadian are not evergreen rainforest. More accurately, chim-
cycle in their beds.) Hernandez-Aguilar (2009) panzees subsist in a range of ecotypes, from
found 5354 nests over a 20 month period at Issa, an woodland savannah (not steppe) to rainforest, with
open-country, savannah area in western Tanzania. mean annual rainfall that range from about 800
These shelters were highly clumped on woodland hill- to more than 2000 mm per year. Many of these
sides, in particular sites that were re-used over and landscapes are vegetationally heterogenous, and chim-
over again. The chimpanzees’ ranging and consequent panzee use of this array of habitat types varies greatly.
nest distribution varied predictably over wet and dry At the other extreme, chimpanzees (unlike baboons,
seasons, reflecting an annual cycle of movement that Papio spp.) do not survive in places that lack surface
reflects availability of surface water and ripe fruit. water for drinking or that lack the riverine forests
However prominent a part these shelters play in their that follow these watercourses, although only a tiny
daily lives, these constructions later will be archaeolo- fraction of such gallery forest will suffice. Copeland’s
gically invisible, being made entirely of woody (2007, 2009) detailed comparison of several open
vegetation. and arid African habitats shows that landscapes with
At the same time, studies of individual nests and annual rainfall in the 500 – 750 mm range cannot sup-
their making have yielded insights: port chimpanzees. Early hominins apparently relied on
Koops et al. (2007) showed that at Nimba, surpris- eating C4 plants and USOs, both of which have yet to
ingly many nests were built on the ground. From be shown to be important in the diets of chimpanzees,
the patterning and size of nests, they hypothesized despite recent prominent findings (Hernandez-Aguilar
that this reflected a pattern of male overnight mate- et al. 2007). When drinking water runs short, that is,
guarding, that is, when an oestrous female nested in during the dry season when water table drops below
a tree, a male seemed to nest on the ground at its the surface, chimpanzees turn to digging wells when
base, to sequester her from the nocturnal attention riverbeds are sandy enough to allow this (Hunt &
of other males. Various functions for nests have been McGrew 2002). Although the wells are dug by hand,
proposed: anti-predator, anti-parasite, anti-disease leaf sponges are used to extract water from the wells;
vector, thermoregulation, etc., but there has yet it would not be surprising to find digging tools used
been no comprehensive study of these hypotheses. to dig wells in other substrates, e.g. mud, gravel, etc.
Meanwhile, Stewart et al. (2007) studied the proximal On a day-to-day basis, chimpanzees must find
characteristics of nests, in terms of their architecture ephemeral food. Frugivores in particular must find
and materials. First-hand empirical data showed that and monitor clumps of food that should be eaten at
chimpanzees prefer comfortable nests, presumably to peak ripeness and which varies from year to year in
gain restorative sleep for their big brains. availability. The same grove that yielded a bumper
The species’ name for the chimpanzee implies a crop last year may not fruit at all this year. The biodi-
cave dweller, yet until recently, there was no record verse array of trees, shrubs and lianas, much less non-
of chimpanzees using caves as shelter. Pruetz (2007) woody plants, may present a potential cornucopia of
reported that the Fongoli chimpanzees, who occupy food, but the daily challenge is how to be in the right
one of the hottest and driest areas in the species’ distri- place at the right time. Various hypotheses have been
bution, regularly use a cave during the hottest season put forward as to how chimpanzees achieve this, but
of the year. They retreat to its cooler environment the strategy turns out to be simple:
during the heat of the day for ‘siestas’ and picnics; Normand et al. (2009) showed that chimpanzees in
overnight, they sleep in arboreal nests, just like other the Taı̈ Forest memorize the locations of thousands of
great ape populations. individual trees. Modelling of the apes’ powerful
Chimpanzees are notoriously hydrophobic, as they spatial memory allows for their ‘rules’ of foraging to
do not swim, which makes watercourses notable bar- be inferred, e.g. travel longer distances to resources
riers to their geographical distribution. However, they that allow longer feeding bouts, revisit more often
enter surface water in certain circumstances: at Fongoli, sources where you last ate for long periods.
they immerse themselves in temporary rain-filled pools But how to acquire such information? Murray et al.
at the beginning of the rainy season, when it is still hot (2008) showed at Gombe that even in adulthood and
and humid; there they rest, groom and play (Pruetz & long after their mothers have died, males return to
Bertolani 2009). Thus, water becomes a thermoregula- the core ranges used by their mothers, especially in
tory device, even when potentially risky. lean times. Resource locations learned during depen-
dent infancy are harvested lifelong.
It is all very well to know what resources are in the
(d) Ranging home range, but how to know where they are, that is,
Although some authors (e.g. Lovejoy 2009) stub- how to navigate optimally between them? Again, var-
bornly continue to characterize evergreen rainforest ious hypotheses have been proposed, e.g. spatial
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3272 W. C. McGrew Chimpanzees and the last common ancestor

orientation by means of landmarks. Normand & become hammers, as they are modified by use
Boesch (2009) show from data on travel directions (Carvalho et al. 2009). That is, tools may change
and distances that Taı̈ chimpanzees have sophisticated functional categories. (Studies of refitting may
mental maps, that is, cognitive two-dimensional help to distinguish reduction products from tool
representations of the landscape that allow them to sets; e.g. Delagnes & Roche 2005). Moreover,
travel from resource to resource in straight lines. application of knowledge from ape tool sets may
help make sense of patterned heterogeneity in
archaeological assemblages, as revealed by multi-
2. DISCUSSION variate statistical analyses.
What can now be said about the LCA, based on what — Composite tools probably were used by the LCA,
has been learned over the past 5 years from field but the challenge is to recognize such combinations
studies of wild chimpanzees? in recovered lithic assemblages. It is not always
Technology is the obvious starting point: clear what was the goal of reduction sequences in
knapping, such as core or flake. The best candidate
— Given the large and varied tool kits of the chimpan- still may be pounding technology, as it seems likely
zee, we can expect that of the LCA to be similar. that flaked stone did not spring de novo with the
That is, tools were made and used not just for Oldowan, but more probably evolved from earlier
food acquisition and processing, but also in self- lithic percussion for other reasons. Perhaps the
maintenance and shelter, as well as in social and analogues to chimpanzee hammers and anvils are
sexual life (not covered here). However, just as there to be found in deposits older than 2.6 Ma?
the size of tool repertoire in chimpanzees is a func- Primatologists should be able to help in seeking
tion of research effort, so it will be in recovering the the pre-Oldowan (Haslam et al. 2009), based on
material culture of the LCA. reliably recognized modifications from chimpanzee
— Most of the presumed technology of the LCA is hammers and anvils. This may help to clarify per-
archaeologically unrecoverable, given its perish- sisting confusion and controversy (e.g. Mora & de
able, organic nature; thus the archaeological la Torre (2005) versus Diez-Martin et al. (2009))
record is biased towards lithics. Short of a time among archaeologists.
machine, this problem is insoluble, but aspects of — Apart from their nest-building, chimpanzees have
chimpanzee behaviour that are universal, such as few compound artefacts. In the evolution of
bed-making or leaf-sponging, are hard to deny to human elementary technology, much is made of
the LCA. the first evidence of hafted weapons, that is, a com-
— As with chimpanzees, the material culture of the pound tool of shaft, point and fixative. However,
LCA will show inter- and intra-regional differences arguably, the earliest known compound technology
(e.g. Schoening et al. 2008). Just as nut-cracking was necklaces of snail shells, as found in Blombos
differs between East and West Africa (Morgan & Cave, South Africa (Henshilwood et al. 2004).
Abwe 2006), despite the common presence of Whether or not the LCA had compound tools is
both prey and raw materials (McGrew et al. unclear, especially as not all components survive
1997), so it is for the LCA. Similarly, just as extrac- equally well, e.g. the spear’s shaft versus its point,
tive foraging for social insects is central to the necklace’s string versus its shells.
Tanzanian populations of chimpanzees, but is lar- — Studies of the acquisition and development of
gely absent in the neighbouring country of chimpanzee technology remind us that some pro-
Uganda, so we should not be surprised to find portion of what is found archaeologically is
such differences in e.g. Kenyan and Ethiopian probably the immature version of the polished
populations of a species of hominin. adult form of material culture. How much debitage
— Subsistence technology in chimpanzees involves reflects ‘honest’ mistakes by youthful learners
reuse of artefacts, whether these are nut-cracking versus clumsy or misguided efforts by adults?
hammers or ant-dipping wands. Especially given This problem probably applies as well to the
that the extent of reuse seems to be a function LCA. Actualistic studies of children of various
of availability of raw materials (and some African ages learning to knap stone might be useful.
forests afford no surface stones bigger than a — Finally, we must repeatedly remind ourselves that
walnut, e.g. Lui Kotale, W. C. McGrew & the LCA was almost certainly not a chimpanzee,
L. F. Marchant 2006, unpublished data), the and vice versa. Just as living apes continue to
same is expected of the LCA. Just as at Bossou, reveal new kinds of technology, so should we
reuse of stone tools may increase the probability expect the same from the LCA. If chimpanzees
of predictable fracture or amplified use – wear that turn out not to use tools to make other tools, or
would leave archaeological signatures in the result- lack important but basic material cultural items
ing artefacts. Lack of data on curation of tools by like the container, or do not transport objects
apes in nature may reflect lack of precise study, as over long distances, we may have found important
evidence exists of such premeditated storage in hominin watersheds (cf. Wynn & McGrew 1989).
captivity (Osvath 2009).
— Given tool sets in chimpanzees, we should expect Regarding diet:
the same in the LCA. But how to recognize
sequential use from a static assemblage? This is — Chimpanzee opportunistic omnivory is clear, and
further complicated by findings that anvils may so it is probably in the LCA. The same inference
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Chimpanzees and the last common ancestor W. C. McGrew 3273

derives from increasing evidence of dietary overlap most chimpanzee field sites do not offer caves,
(e.g. monkey-hunting) between chimpanzee and although this has never been systematically studied.
bonobo, although important differences remain — We now know that chimpanzee nests are more
between these taxa (e.g. extractive foraging for complex structures than hitherto realized, and
insects). this may imply that beyond a certain point of
— Recent findings of chimpanzee use of USOs para- investment of time and effort, they began to be
doxically show apes to be capable of harvesting reused. This raises the possibility of home bases,
these foodstuffs, yet in no known population are already hinted at in the non-random distribution
they a staple (cf. Hockings et al. 2009, for data of chimpanzee nest sites on the landscape. But
on USOs as fallback foods). Experimental studies until we know the fitness-enhancing function of
need to be done on the limits of chimpanzee- beds, it would be rash to infer the same for the
digging. Similarly, chimpanzees commonly LCA. Anti-predation is assumed, but equally
consume the pith of C4 plants, yet not the seeds attractive alternative hypotheses are there to be
or corms, and so their stable isotope data are con- tested. The presence of ground nests is sometimes
fusing (Sponheimer et al. 2006). (It seems likely presumed to be based on local release from preda-
that staple exploitation of cereals requires grinding tion, but no correlative study of sympatric large
technology, which seems to be absent in wild chim- carnivores and apes has been done.
panzees, but apparently has not been tested with
apes in captivity.) Or, it may be that profitable On ranging and foraging:
use of USOs and cereals requires treatment by
fire, that is, cooking, which came much later — Chimpanzees are nomadic over areas that can be
in human evolution (Carmody & Wrangham large, that is, tens or even hundreds of square kilo-
2009). Here, studies of wild chimpanzees are not metres. If the singlemost obvious influence on this
yet helpful in hypothesizing about the LCA. ranging is food availability, the more crucial limit-
— Chimpanzees are wide-ranging foragers, and their ing factors may be drinking water and cover.
patterns of ranging map onto the distribution of Well-digging, especially with the technological
their resources, as in any other organism. What assistance of digging tools and containers, appears
we now are beginning to know is the extent of to allow an expanded ecological niche. (Unlike
their intelligent foraging, and it exceeds our expec- temperature or humidity, which turn out not to
tations, e.g. about spatial memory. This upgrades be so important.) Similarly, no matter how dry
our estimation of the LCA, but inferring the and open the eco-type inhabited, every known
timing and spacing of resources in the archaeologi- population of great apes seems to require access
cal record is problematic. to trees for shelter construction. Even savannah-
— Recent findings on chimpanzee hunting confirm its dwelling chimpanzees need their ribbons of gallery
seductiveness for evolutionary scenarios. (Conver- forest. The same was probably true of the LCA.
sely, scavenging’s role seems less and less
In conclusion, even if one-tenth of what has been
important, at least until after the LCA, in the
learned in the last five years about wild chimpanzees
hominin lineage.) However, estimations of the
is applicable to the LCA of living apes and humans,
importance of hunting, based on chimpanzees,
then the case has been made for preserving them.
must be tempered: Most chimpanzee hunting is
Referential modelling requires living proxies upon
done arboreally, by ‘four-handed’ hunters who
which to base the models, and current expectations
can leap about in the canopy, pursuing monkeys.
are that wild populations of great apes may be gone
This is not likely to be instructive about hunting
by the middle of the current century. Both primatolo-
by terrestrial bipeds, even if it applies to the
gists and palaeoanthropologists should work together
LCA, who may have practised ambush hunting
to save them.
on the ground, as well as pursuit hunting in the
treetops. More significantly, the function of carniv- Most of the author’s data and ideas were supported by the
ory is revealed to be much richer than expected: US National Science Foundation, Researching Hominid
sharing meat may drive social and sexual life, Origins Initiative (Award no. BCS-0321893) grant awarded
to Tim White and to the late Clark Howell. I am grateful
almost as a currency (although many of the same to S. Carvalho, L. F. Marchant, P. Mellars and A. Walker
arguments probably apply also to honey). for comments on the manuscript.

On shelter:
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