Body Size, Body Shape, and the Circumscription of the Genus Homo

Author(s): Trenton W. Holliday

Source: Current Anthropology, Vol. 53, No. S6, Human Biology and the Origins of Homo
(December 2012), pp. S330-S345
Published by: The University of Chicago Press on behalf of Wenner-Gren Foundation for
Anthropological Research
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 S330 Current Anthropology Volume 53, Supplement 6, December 2012

Body Size, Body Shape, and the

Circumscription of the
Genus Homo
by Trenton W. Holliday

Since the 1984 discovery of the Nariokotome Homo erectus/Homo ergaster skeleton, it has been almost axiomatic
that the emergence of Homo (sensu stricto) was characterized by an increase in body size to the modern human
condition and an autapomorphic shift in body proportions to those found today. This was linked to a behavioral
shift toward more intensive carnivory and wider ranging in the genus Homo. Recent fossil discoveries and reanalysis
of the Nariokotome skeleton suggest a more complex evolutionary pattern. While early Homo tend to be larger
than Australopithecus/Paranthropus, they were shorter on average than people today. Reanalysis of the Nariokotome
pelvis along with the discovery of additional early and middle Pleistocene pelves indicate that a narrow bi-iliac
(pelvic) breadth is an autapomorphy specific to Homo sapiens. Likewise, it appears that at least some early Homo
(even those referred to H. ergaster/H. erectus) were characterized by higher humero-femoral indices than the H.
sapiens average. All these data suggest a pattern of mosaic postcranial evolution in Homo with implications for the
increased ranging/carnivory model of the origin of Homo as well as for which species are included within the Homo
hypodigm.

Introduction
The oldest and most persistent questions in paleoanthropol-
ogy are those that pertain directly to us. Specifically, the two
longest-lived questions in our discipline are (1) the origin of
our own species, Homo sapiens, which occurred sometime
around the end of the middle Pleistocene; and (2) the origin
of our own genus, Homo, which most likely occurred in the
late Pliocene. The questions surrounding the origins of these
taxa are not merely of phylogenetic bookkeeping interest. Be-
cause they involve our direct ancestors, we tend to imbue
these questions with a suite of functional/adaptive explana-
tions (e.g., intelligence and/or spoken language in the case of
modern human origins and increased encephalization, larger
body and home-range size, more efficient bipedality, and in-
creased carnivory in the case of the origins of the genus
Homo). In light of humanitys global distribution, invoking
adaptation when our own species/genuss evolution is con-
cerned is certainly warranted. Our widespread geographic dis-
tribution is even more impressive when one considers that
among the African hominids (sensu lato), humans are evo-
Trenton W. Holliday is Professor of Anthropology at Tulane
University (101 Dinwiddie Hall, 6823 St. Charles Avenue, New
Orleans, Louisiana 70118, U.S.A. [thollid@tulane.edu]). This paper
was submitted 12 XII 11, accepted 11 VI 12, and electronically
published 14 IX 12.
lutionarily the only successful taxon, because the other African
apes appear poised for extinction in the wild, and importantly,
their decline is not likely merely the result of recent habitat
destruction due to an ever-expanding human population but
rather may have more to do with their overreliance on a highly
K-selected reproductive strategy coupled with a decreased
population growth rate associated with larger brains (Isler
and van Schaik 2012; Lovejoy 1981, 2009).
This paper studies early (ca. 2.01.5 million years ago
[mya]) Homo specifically with regard to its postcranial skel-
eton, body size, and limb and body proportions. The study
of limb and body proportions of early Homo in particular
has led to extensive debate as to whether the earliest members
of the genus had limb and body proportions similar to those
of Australopithecus (who are presumed to have included more
of an arboreal component in their locomotor behavior) or,
in contrast, whether their limb and body proportions were
more similar to those of modern humans, which could be
taken as indicative of fully committed terrestrial bipedality
(Bramble and Lieberman 2004; Haeusler and McHenry 2004,
2007; Hartwig-Scherer and Martin 1991; Holliday and Fran-
ciscus 2009; McHenry and Berger 1998a, 1998b; Pontzer 2012;
Pontzer et al. 2010; Reno et al. 2005; Richmond, Aiello, and
Wood 2002). Observed (or presumed) differences in body
size and proportions between Homo sapiens and some early
Pleistocene hominins referred to Homo have even played a
role in some researchers call to remove key specimens (and

2012 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2012/53S6-0006$10.00. DOI: 10.1086/667360

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Holliday Body Size and Shape in Early Homo
even presumed species-level taxa) from the genus Homo al-
together (Collard and Wood 2007; Wood and Collard 1999a,
1999b).
Circumscribing the Genus Homo
In order to discuss body size and shape in the genus Homo,
it is critical to determine the composition of that genus. In
zoology, there are generally four nonmutually exclusive sets
of criteria by which species taxa are considered congeneric:
(1) recency of common ancestry, (2) ecological/adaptive sim-
ilarity, (3) genetic divergence, or (4) morphological similar-
ities. However, there is little agreement among zoologists as
to how exactly this is done, and, as recently pointed out by
Collard and Wood (2007), relatively little attention has been
paid by taxonomists as to how the genus category is opera-
tionally defined. Ernst Mayr (1942:283), one of the founders
of the school of evolutionary taxonomy, defined a genus as
one species or a group of species of presumably common
phylogenetic origin, separated by a decided gap from other
similar groups. Mayrs definition therefore allows for grade
characters (or at a minimum, perceived gaps between them)
to be involved in circumscribing taxa, making it unacceptable
to strict practitioners of phylogenetic systematics (cladistics)
as developed by Willi Hennig.
Hennig (1966) argued that all biological taxa be strictly
monophyletic (what Mayr [2000] suggests should instead be
referred to as holophyletic); that is, taxa at all levels should
be made up of all the descendents of an ancestral taxon re-
gardless of their plesiomorphic versus apomorphic (i.e.,
grade) status. Of course, the criterion of monophyly alone
does not help one distinguish a monophyletic genus from a
monophyletic tribe, subfamily, or family. Hennig (1966)
therefore suggested that time since divergence be the second
major criterion for circumscribing genera, but he also argued
that rather than using a universal one size fits all time frame
(i.e., all genera everywhere should be 810 million years old),
temporal criteria should be specific to each particular bio-
logical group studied. While admittedly subjective, this would
seem appropriate given vast differences in divergence times
among long-recognized good zoological higher taxa. For
example, molecular data suggest that the two genera of Asiatic
horseshoe crabs (Tachypleus and Carcinoscorpius) diverged
from their North American cousins (genus Limulus) ca. 45
60 mya (Avise, Nelson, and Sugita 1994), which may be earlier
than the divergence of the Old World anthropoids (infraorder
Catarrhini) from the New World monkeys (infraorder Pla-
tyrrhini) and certainly millions of years earlier than diver-
gence of the family Hominidae (apes and humans) from the
Cercopithecidae (Old World monkeys).
Thus, it is evident that neither the phylogenetic systematic
description of genera nor the definition of genera employed
by evolutionary taxonomists is without its flaws, but it should
be possible to combine the strengths of both into a better
means by which to define generic taxa. Applying this logic,
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S331
Wood and Collard (1999a, 1999b) used an operational defi-
nition of the genus category that combined what they viewed
as the stronger aspects of both Mayrs (1942) and Hennigs
(1966) approaches. They recognized that a strict Hennigian
approach to defining genera gave no real guidance as to how
to circumscribe genera versus other higher Linnaean cate-
gories and that Mayrs (1942) approach was faulty in that it
could lead to the recognition of paraphyletic generataxa
that are biologically unreal in that they do not reflect the
true evolutionary relationships of the taxa in question. In-
stead, Wood and Collard (1999b:66) define a genus as a
species, or monophylum, whose members occupy a single
adaptive zone. Importantly, their definition of a genus does
not restrict any particular adaptive zone to a single genus,
instead allowing for the possibility that species assigned to
different genera will occupy the same adaptive zone, but it
does prevent species in the same genus from occupying dif-
ferent adaptive zones (Collard and Wood 2007:1584). An
adaptive zone is characterized by Wood and Collard (1999a:
202) as being related to an organisms phenotype and its
ability to maintain homeostasis, acquire food, and produce
offspring. While recognizing that not all of these aspects of
the phenotype are easily recognizable in the fossil record,
Wood and Collard (1999a, 1999b) maintain that certain fea-
tures, such as the size of the masticatory apparatus, relative
brain size, ontogenetic pattern, body size and shape, and lo-
comotor behavior all leave phenotypic traces observable in
hominin fossils.
In circumscribing the genus Homo, then, Wood and Collard
(1999a, 1999b) argued that two main criteria must be met:
(1) that the genus Homo be monophyletic (holophyletic) and
(2) that all its members share a common adaptive strategy or
zone. To test the first criterion, Wood and Collard (1999a,
1999b) reported the results of multiple cladistic analyses, some
of which failed to consistently link Homo habilis and Homo
rudolfensis as sister taxa to Homo sapiens to the exclusion of
the australopiths.
For the second (i.e., the adaptive-zone) criterion, Wood
and Collard evaluated whether a presumed member species
of the genus Homo was more similar to the type species of
the genus Homo (H. sapiens) or Australopithecus (Australo-
pithecus africanus) for the following adaptive complexes: (1)
body size, (2) body shape, (3) locomotion, (4) mastication,
(5) growth and development, and (6) relative brain size.1 They
found that Homo ergaster, Homo erectus, Homo heidelbergensis,
and Homo neanderthalensis were more similar to H. sapiens
than A. africanus for all, or at least the vast majority, of these
adaptive complexes (the clade including these five species will
1. Anton (2012) points out that comparing 2.01.5-million-year-old
fossil hominins to modern humans (Homo sapiens) in order to refer them
to Homo (or not) is problematic in that it will tend to disregard similarities
shared between Homo habilis/Homo rudolfensis and Homo ergaster/Homo
erectus, instead emphasizing differences between earliest Homo and the
most derived taxon in the clade (H. sapiens), a taxon that also happens
to be far removed from early Homo in time.
 S332
here be referred to as Homo [sensu stricto]). In contrast,
according to their analyses, H. rudolfensis and H. habilis were
decidedly more Australopithecus-like. Wood and Collard
therefore came to the conclusion that these two species failed
both the phylogenetic and the adaptive-zone criteria for in-
clusion in the genus Homo and suggested that they instead
be referred to the genus Australopithecus.
More recently, Collard and Wood (2007) redid their 1999
analyses taking into account new fossil data as well as more
recently published cladistic analyses. As before, the cladistic
analyses remained somewhat inconsistent in clustering H. ha-
bilis and H. rudolfensis with later Homo to the exclusion of
other taxa. Based on new data, they did find some changes
in adaptive-zone patterning from their 1999 results (partic-
ularly with regard to the pattern of growth and development
of H. ergaster; and see Schwartz 2012); however, their ultimate
conclusion remained that neither H. habilis nor H. rudolfensis
should be included in the Homo hypodigm (Collard and
Wood 2007).
This paper evaluates five reflections of body size and shape
in early Homo (sensu lato) that figure into the adaptive-zone
half of the circumscription of the genus Homo. The features
evaluated here allow the examination of three of the six adap-
tive complexes as defined by Wood and Collard (1999a,
1999b): (1) body size, (2) body shape, and (3) locomotion.
The current analyses also include recently published data not
available to Wood and Collard (1999a, 1999b) or Collard and
Wood (2007). This paper will attempt to assess whether (1)
the emergence of the genus Homo is associated with an in-
crease in locomotor efficiency specifically as related to ter-
restrial bipedality and its relationship to ranging behavior,
and (2) H. habilis and H. rudolfensis should be removed from
the genus Homo based on Wood and Collards adaptive-zone
criteria for body size and shape and locomotion. The five
specific postcranial features to be examined are (1) relative
lower limb length, (2) humero-femoral proportions, (3) rel-
ative forearm length, (4) relative pelvic breadth, and (5) body
size as reflected in predicted body mass and a proxy for stat-
ure.
Relative Lower Limb Length
Longer limbs are known to increase the efficiency of animal
locomotion by reducing the number of strides necessary to
cover any given distance (Jungers 1982; Pontzer 2007, 2012;
Steudel-Numbers 2006; Steudel-Numbers and Tilkens 2004).
In this light, it has long been argued that Australopithecus had
relatively short lower limbs while Homo (sensu stricto) was
characterized by longer lower limbs (Bramble and Lieberman
2004; Jungers 1982, 1991a; Jungers and Stern 1983; Pontzer
et al. 2010; Richmond, Aiello, and Wood 2002). However,
there are many recent data that challenge this assumption.
First, this view, while prevalent, has been largely based on the
diminutive A.L. 288-1 Australopithecus afarensis (Lucy)
specimen (Jungers 1982, 1991a; Jungers and Stern 1983). The
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Current Anthropology Volume 53, Supplement 6, December 2012
problem with this line of evidence is that lower limb length
shows positive allometry in humans (Auerbach and Sylvester
2011; Holliday and Franciscus 2009; and see Pontzer 2012)
such that the expectation is for small-bodied hominins such
as Lucy, including members of the genus Homo, to have
shorter lower limbs (Franciscus and Holliday 1992; Holliday
and Franciscus 2009; but see Haeusler and McHenry 2004).
The discovery of diminutive late Pleistocene (but clearly genus
Homo) fossils at the site of Liang Bua on Flores (referred to
the species Homo floresiensis; Brown et al. 2004), which are
characterized by relatively short lower limbs (Jungers et al.
2009), provides further support for this idea no matter what
those specimens ultimate taxonomic status within our genus
(Holliday and Franciscus 2009).
Likewise, there have been numerous recent discoveries and/
or reanalyses of larger-bodied members of the genus Aus-
tralopithecus who (unlike Lucy) are characterized by rela-
tively elongated lower limbs. These include the A. afarensis
specimens A.L. 333-3, A.L. 827-1 (Harmon 2005; Kimbel and
Delezene 2009), and KSD-VP-1/1 (Haile-Selassie et al. 2010)
and the (presumed) Australopithecus africanus specimen StW
99 (T. W. Holliday, unpublished data). The overall pattern of
lower limb length relative to size in hominins is illustrated
by the scatterplot in figure 1, which shows femoral length
regressed on femoral head diameter. The former measurement
reflects lower limb length, while the latter reflects overall body
mass. Note that Homo sapiens is largely separated from Pan
and Gorilla in bivariate space in that for any given femoral
head size, our species is expected to have a much longer femur.
With two exceptions, all of the fossil hominins fall within the
95% confidence limits about the H. sapiens individuals for
this relationship and fall outside the 95% confidence limits
about Pan and Gorilla. The exceptions, notably, are the di-
minutive A.L. 288-1 (Lucy) and Liang Bua 1 (holotype of
H. floresiensis) specimens, which fall among the chimpanzees.
The slightly larger (but still diminutive by modern human
standards) A.L. 152-2 A. afarensis individual falls at the limits
of the H. sapiens sample and just beyond the 95% confidence
limits about the chimpanzee individuals. Note, however, that
this individual falls almost directly on a modern human
Pygmy individuals values for both measurements. In con-
trast, the larger-bodied australopiths (A.L. 333-3, A.L. 827-1,
KSD-VP-1/1, and StW 99), while characterized by smaller
femoral heads than most members of the genus Homo (long
known to be an australopith characteristic; Galik et al. 2004;
Harmon 2009; Jungers 1988, 1991a; Kennedy 1983; Lovejoy,
Heiple, and Burstein 1973; Napier 1964; Richmond and Jung-
ers 2008; Robinson 1972; Ruff 1988; Walker 1973), nonethe-
less fall among the H. sapiens sample for this relationship and
beyond the confidence limits about the African ape samples,
as do the early Homo specimens KNM-ER 1472 and 1481,
KNM-WT 15000 (the Nariokotome Boy), and Dmanisi
4507.
A scaling difference in femoral head size between Homo
and the australopiths has also been documented (Jungers
Holliday Body Size and Shape in Early Homo S333

Figure 1. Scatterplot of femoral length regressed on femoral head diameter for Pleistocene/Holocene Homo sapiens, Pan, Gorilla,
and Pliocene/Pleistocene early hominins (and Liang Bua 1). Australopiths are indicated by open squares, fossil Homo by filled
squares. The ordinary least squares regression lines for the comparative samples are represented by solid lines with the 95% confidence
limits for the individuals indicated about them. The reduced major axis (RMA) regression lines for the comparative samples are
the dashed lines. Homo sapiens RMA formula: y p 8.074x 77.175 ; Pan RMA formula: y p 7.159x 56.2 ; Gorilla RMA formula:
y p 6.439x 53.28.

1988; Ruff 1990). It is probable that this difference is in some
way related to a shift in locomotor repertoire between Homo
and the australopiths, but the exact nature of such a shift is
uncertain, especially when one considers that there are ap-
parent differences in upper versus lower limb articular and
length proportions even within the genus Australopithecus (see
below). Despite this caveat, the pattern of relative lower limb
length revealed in figure 1 is mirrored in plots for which body
mass and not femoral head diameter is plotted along the X-
axis (Franciscus and Holliday 1992; Holliday and Franciscus
2009; Pontzer 2012; Pontzer et al. 2010).
Humero-Femoral Proportions
The discovery of the A.L. 288-1 specimen, with its nearly
intact humerus and femur, led to the confirmation that Aus-
tralopithecus afarensis was characterized by very different
humero-femoral proportions than the genus Homo (or, for
that matter, the genus Pan), with longer humeri relative to
their femora than Homo and much shorter humeri relative
to their femora than Pan (Johanson et al. 1982). What fol-
lowed were years of intense debate as to whether the humero-
femoral proportions of Lucy were due to a long humerus,
a short femur, or a combination of both (Franciscus and
Holliday 1992; Jungers 1982, 1991a; Jungers and Stern 1983;
Wolpoff 1983). It now seems probable that the last of these
alternative explanations best accounts for the data (Holliday
and Franciscus 2012).
It has also been argued that the humero-femoral propor-
tions of Australopithecus africanus and/or Homo habilis were
as different, or more different, from modern humans than
those of A. afarensis (Hartwig-Scherer and Martin 1991; Mc-
Henry and Berger 1998a). Unfortunately, in the case of H.
habilis, at least, this supposition is based on faulty data
specifically the femoral length of the OH 62 specimen, which
cannot be reliably reconstructed with any degree of confidence
(see below). Another presumed H. habilis partial skeleton,
KNM-ER 3735, is even less complete (Haeusler and McHenry
2007). Similarly, there is a relative dearth of associated A.
africanus skeletons, and those that are available (e.g., Sts 14
and StW 431) lack complete limb bones.

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 S334 Current Anthropology Volume 53, Supplement 6, December 2012

Patterning in humero-femoral proportions among African
hominids (sensu lato) is shown in figure 2, which is a scat-
terplot of humeral length regressed on femoral length. As has
long been appreciated, gorillas have the longest humeri rel-
ative to their femora, followed by chimpanzees, while humans
have the shortest humeri relative to the length of their femora
(or the longest femora relative to their humeri). The first
pattern evident in the graph is that the humero-femoral pro-
portions of OH 62 are impossible to assess (as was argued
by Haeusler and McHenry 2004, 2007; Korey 1990; Reno et
al. 2005; Richmond, Aiello, and Wood 2002). The primary
problem in the assessment of OH 62s humero-femoral pro-
portions is unlikely to be its humerus; Haeusler and McHenry
(2004:446) point out that the OH 62 humerus is relatively
well preserved, lacking only the proximal and distal extrem-
ities. Its length can be estimated with only minor error. They
estimate humeral length of the specimen at 270 mm. Here
an estimate of 264 mm from Johanson et al. (1987) is used
(these two estimates differ from each other by only 2%).
Despite the fact that Korey (1990) argues there is a high
degree of variability in humeral length estimation for OH 62,
the more intractable problem with assessing its humero-fem-
oral proportions lies with its femur. As noted by Reno et al.
(2005), it is likely that less than half of the femurs length is
preserved, and among hominids (sensu lato) there is no pre-
dictable relationship between proximal femoral length and
maximum (or bicondylar) femoral length. Because Johanson
et al. (1987) argued that the OH 62 shaft was smaller and
less robust than that of A.L. 288-1, some researchers (e.g.,
Haeusler and McHenry 2004; Richmond, Aiello, and Wood
2002) have taken to using Lucys femoral length of 280 mm
as the minimum length for the OH 62 femur, a convention
followed here. Haeusler and McHenry (2004) argue, however,
that the longer (albeit damaged; see Green, Gordon, and Rich-
mond 2007) OH 34 femur is a better analog for the OH 62
femur than is A.L. 288-1. They therefore used their estimated
OH 34 femoral length of 374 mm as the maximum (potential)
femoral length for OH 62; the same is done here. This equates
with nearly 100 mm of difference between the minimum and
maximum estimates of the OH 62 femoral length. Thus, even
in the absence of error in humeral length, we can say nothing
about OH 62s humero-femoral proportions because, as is
evident in figure 2, they could range from somewhat more
chimpanzee-like than Lucy (as was maintained by Hartwig-

Figure 2. Scatterplot of humeral length regressed on femoral length for Pleistocene/Holocene Homo sapiens, Pan, Gorilla, and
Pliocene/Pleistocene early hominins (and Liang Bua 1). Australopiths are indicated by open squares, fossil Homo by filled squares.
The ordinary least squares regression lines for the comparative samples are represented by solid lines with the 95% confidence limits
for the individuals indicated about them. The reduced major axis (RMA) regression lines for the comparative samples are the dashed
lines. Homo sapiens RMA formula: y p 0.748x 14.432; Pan RMA formula: y p 1.01x 4.15; Gorilla RMA formula: y p
1.092x 26.715.

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Holliday Body Size and Shape in Early Homo S335

Scherer and Martin 1991) to falling almost directly on the
Homo sapiens reduced major axis (RMA) and ordinary least
squares (OLS) regression lines.
Note, too, that as was the case for the previous relationship,
the only fossil hominins to fall squarely outside of the H.
sapiens sample in figure 2 are the diminutive A.L. 288-1 and
LB1 specimens. The limb-segment lengths of the 2.5-million-
year-old taxonomically unassigned remains from Bouri
(BOU-VP-12/1) were estimated anatomically by Asfaw et al.
(1999) at 226236 mm for the humerus and 335348 mm
for the femur, which places them squarely among H. sapiens
in bivariate space (although there is likely a problem with
these estimates; see below). The early Pleistocene Homo re-
mains from Dmanisi and KNM-WT 15000 also fall among
the more recent humans, although Dmanisi 4167/4507 falls
on the 95% confidence limits about the recent human in-
dividuals. This graphic result is, however, somewhat different
from that depicted in figure 3b of Lordkipanidze et al. (2007),
where Dmanisi 4167/4507 appears to fall in the middle of the
recent human scatter for the humerus length to femoral length
bivariate relationship. On closer inspection of Lordkipanidze
et al. (2007), however, it becomes evident that the 295-mm
maximum humeral length of Dmanisi 4507 is erroneously
represented as ca. 275 mm in their figure 3b.
A histogram of humero-femoral indices of the H. sapiens
sample is presented in figure 3 with the positions of the fossils
that preserve both a humerus and a femur indicated by ar-
rows. As has been noted multiple times (e.g., Brown et al.
2004; Johanson et al. 1982; Jungers 1982; McHenry 1978),
A.L. 288-1 and LB1 lie far (5.6 and 6.6 standard deviations,
respectively) above the Homo sapiens mean. While falling
within the anatomically modern human range, Dmanisi 4507
and KNM-WT 15000 still have high index values because they
lie at the modern human 97th and 84th percentiles, respec-
tively. In fact, a special case of Students t-test for comparing
a single individual to a sample (Sokal and Rohlf 1981) finds
that Dmanisi 4507 is statistically significantly different (at
P ! .05) from modern humans (ts p 2.134, P p .033),
whereas KNM-WT 15000 is not (ts p 1.014, P p .311). Given
that Dmanisi dates to ca. 1.77 mya (Gabunia, Vekua, and
Lordkipanidze 2000) and KNM-WT 15000 dates to ca. 1.53
mya (Brown and McDougall 1993), if each is reflective of its
population, and assuming that these skeletons represent a
lineage in the broadest sense, then it may be that humero-
femoral indices decreased through time in early Homo.
In light of the above patterning, the extremely low humero-
femoral index values (ranging from below the first to the 32nd
percentiles of H. sapiens) of the 2.5-million-year-old BOU-
VP-12/1 specimen appear particularly incongruous. This
could be due to the humerus and femur coming from different
individuals or (more likely) one or both of the estimated long-
bone lengths being erroneously reconstructed. If the latter is
the case, the most probable scenario, given the preservation
of the remains shown in figure 4 of Asfaw et al. (1999), is
that the humeral length of the specimen has been underpre-
dicted, because the femur preserves the diaphysis from the
inferior margin of the neck to an area just proximal to the
lateral epicondyles, and therefore its length should be accu-
rately reconstructed. In contrast, while on the humerus a
proximal portion of the medial epicondyle is preserved (giving

Figure 3. Histogram of humero-femoral indices, Pleistocene/Holocene Homo sapiens sample with the values (or in the case of BOU-
VP-12/1, the range of potential values) of fossil hominins indicated by arrows. Homo sapiens mean p 71.5; SD p 2.3.

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 S336 Current Anthropology Volume 53, Supplement 6, December 2012

one a solid idea of where the distal end of the bone should
be), it is impossible to ascertain from the figure alone how
much of the proximal diaphysis is present.
Relative Forearm Length
One of the key features long thought to distinguish the aus-
tralopiths from the genus Homo is that the former are said
to have had longer (and therefore more apelike) forearms
than the latter (Asfaw et al. 1999; Haeusler and McHenry
2004; Howell and Wood 1974; Kimbel, Johanson, and Rak
1994; McHenry 1978; but see Drapeau and Ward 2007; Haeus-
ler and McHenry 2007). A scatterplot of radius length re-
gressed on femoral length is shown in figure 4. As expected,
for any given femoral length, humans have much shorter radii
than the African apes. Note, too, that for this relationship,
gorilla radii appear to lie along a continuation of the chim-
panzee trajectory, something that was not the case for the
humerus.
With regard to the fossils, the left ulna of A.L. 288-1 is
preserved in two pieces that make only ambiguous contact
with each other. Hausler (2001) believes the bony contact
between these two pieces to be good, and when they are
joined, the ulnar length of 191 mm suggests to him a radius
length of ca. 181 mm. However, Richmond, Aiello, and Wood
(2002) published a humanlike radius length of 174 mm for
Lucy based on Schmids (1983) reconstruction (this value
closely corresponds to the radius length [175 mm] estimated
from a 191-mm ulnar length for a sample of recent humans;
T. W. Holliday, unpublished data). Kimbel, Johanson, and
Rak (1994) generate a radius length of 206 mm for the spec-
imen, and Asfaw et al. (1999) generated a chimpanzee-like
A.L. 288-1 radius length of 215 mm. Here I use this last
estimate as the specimens maximum length and 174 mm as
its minimum. As seen in figure 4, the result is a radius that
perhaps falls just outside the modern human samples 95%
confidence limits but is almost certainly longer than would
be expected for a modern human of its diminutive size. Yet
even the longest length estimate for the specimen does not
fall near the Pan 95% confidence limits, a result consistent
with the findings of Haeusler and McHenry (2007) and
Drapeau and Ward (2007), who found no evidence of chim-

Figure 4. Scatterplot of radius length regressed on femoral length for Pleistocene/Holocene Homo sapiens, Pan, Gorilla, and Pliocene/
Pleistocene early hominins (and Liang Bua 1). Australopiths are indicated by open squares, fossil Homo by filled squares. The
ordinary least squares regression lines for the comparative samples are represented by solid lines with the 95% confidence limits
for the individuals indicated about them. The reduced major axis (RMA) regression lines for the comparative samples are the dashed
lines. Homo sapiens RMA formula: y p 0.623x33.815; Pan RMA formula: y p 1.175x70.195; Gorilla RMA formula: y p
0.93x 3.713.

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Holliday Body Size and Shape in Early Homo
panzee-like antebrachial elongation in Australopithecus afa-
rensis.
As was done for A.L. 288-1, the radius length of LB1 was
estimated from its nearly complete right ulna at 190 mm
(Morwood et al. 2005), placing it near the middle of the
predicted range of A.L. 288-1 and well beyond the 95% con-
fidence limits for both Homo and Pan, albeit closer to the
former than to the latter. The taxonomically unassigned BOU-
VP-12/1 specimen falls beyond the range of both Pan and
Homo for this bivariate relationship, which might be expected,
given its early date. However, recall that at least some of the
estimated limb-segment lengths for this specimen are suspect.
The radius length (255 mm, predicted from ulnar length by
Walker and Leakey [1993]) of KNM-WT 15000 falls just be-
yond the 95% confidence limits about the Homo sapiens in-
dividuals while lying within the scatter of the H. sapiens sam-
ple. The radio-femoral (RL/FL # 100) index for KNM-WT
15000 (59.0) falls at the 98th percentile of the H. sapiens
sample (n p 1,060) and is statistically significantly different
from that sample at P ! .05 (ts p 2.000; P p .0457). While
this result is based on a juvenile (who is potentially patho-
logical; Ohman et al. 2002), it does suggest that relative fore-
arm length may have decreased over time in Homo.
Relative Pelvic Breadth
Narrow pelves have long been known to be a human autap-
omorphy. They have been hypothesized to be related to ob-
stetrical selection (Lovejoy 1988; Simpson et al. 2008), re-
duced gut size (Aiello and Wheeler 1995), or thermal adap-
tation to hot climates (Weaver and Hublin 2009). This last
explanation seems unlikely in light of the discovery of the
tropical yet wide Gona BSN49/P27 pelvis, which has been
attributed to Homo (Simpson et al. 2008; but see Ruff 2010).
It is, however, possible that a narrower pelvis in humans is
related to increasing locomotor efficiency, given that a broad
pelvis is less biomechanically stable and less efficient in bipedal
locomotion (Bramble and Lieberman 2004; Langdon 2005;
Rosenberg and Trevathan 2007), and in light of the fact that
more cursorial members of the Canidae, including dog breeds
such as greyhounds, tend to have narrower pelves than less
cursorial canids (Carrier, Chase, and Lark 2005; Schutz et al.
2009). This hypothesis enjoys little experimental support,
however, because in laboratory settings pelvic breadth is not
associated with locomotor efficiency in humans (Pontzer
2012) and given the fact that effective limb length alone ex-
plains 98% of the variance in locomotor efficiency across a
wide range of terrestrial animals (Pontzer 2007).
A scatterplot of bi-iliac (pelvic) breadth regressed on ace-
tabular height is presented in figure 5. The former is a measure
of body breadth while the latter is a pelvic reflection of body
mass. Note that Homo sapiens and Pan troglodytes have nearly
parallel regression lines, with the chimpanzees differing from
modern humans in that for any given acetabular size they are
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S337
expected to have a wider bi-iliac breadth. Irrespective of their
generic classification, the fossil hominins tend to fall into two
clusters: a small-bodied cluster and a large-bodied cluster. The
three Australopithecus specimens for which bi-iliac breadth
can be reconstructed (A.L. 288-1 [Australopithecus afarensis],
Sts 14 [Australopithecus africanus], and MH2 [Australopithecus
sediba]) fall among the wide-trunked chimpanzees in bivariate
space for this relationship, although Sts 14 and MH2 also fall
just within the 95% confidence limits about the H. sapiens
individuals. The recently described 1.0-million-year-old
BSN49/P27 pelvis from Ethiopia, referred to Homo by Simp-
son et al. (2008), has a wider pelvis than expected for even
a Pan individual of its body size. The Gona pelvis lies closest
in bivariate space to the female Levantine Neanderthal spec-
imen Tabun C1, the bi-iliac breadth of which was acquired
from the virtual reconstruction of that pelvis by Weaver and
Hublin (2009). This specimen also falls well beyond the 95%
confidence limits of the H. sapiens individuals.
The larger-sized fossil hominins on the right side of the
plot include KNM-WT 15000, two middle Pleistocene hom-
inins (Jinnuishan and Atapuerca Pelvis 1), and two Nean-
derthals (La Chapelle-aux-Saints 1 and Kebara 2). The Na-
riokotome specimen falls at the margins of the H. sapiens
sample, but its position in bivariate space is less reliable than
the other fossils because of two factors. The first is that its
acetabular height was estimated from femoral head diameter.
The second is that its predicted adult bi-iliac breadth is es-
timated by Ruff (2010), and there is considerable debate as
to how much further growth would be expected for the spec-
imen (Graves et al. 2010) and even as to how wide the Na-
riokotome pelvis was in its juvenile state (Ruff 1995, 2010;
Simpson et al. 2010). The two middle Pleistocene Homo spec-
imens, Atapuerca Pelvis 1 and the Jinnuishan pelvis, have
extremely wide bi-iliac breadths such that they fall well beyond
the 95% confidence limits about the H. sapiens individuals.
The La Chapelle-aux-Saints 1 bi-iliac breadth used here
(292 mm) is taken from Trinkauss (2011) recent reconstruc-
tion. While Neanderthals are known for their wide bi-iliac
breadths (Churchill 1998; Holliday 1997; Ruff 2002; Steeg-
mann, Cerny, and Holliday 2002), among the large Nean-
derthals included in the plot, La Chapelle-aux-Saints 1 falls
within the H. sapienss scatter and in fact falls below the H.
sapiens RMA line. This indicates that La Chapelle has a re-
markably narrow pelvis for nonmodern Homo. This is in part
due to the specimens remarkably narrow sacrum (falling at
the 15th percentile of the H. sapiens [n p 726] sample). The
specimen also lacks the marked iliac flare seen in Kebara 2.
In contrast, Kebara 2, with a bi-iliac breadth of 313 mm, falls
just beyond the 95% confidence limits of the H. sapiens sam-
ple.
Body Size (Mass and Stature)
It has long been posited that body size increased with the
emergence of the genus Homo. Body size is an important
 S338 Current Anthropology Volume 53, Supplement 6, December 2012

Figure 5. Scatterplot of bi-iliac breadth regressed on acetabular height for Pleistocene/Holocene Homo sapiens, Pan, and Pliocene/
Pleistocene early hominins. Australopiths are indicated by open squares, fossil Homo by filled squares. The ordinary least squares
regression lines for the comparative samples are represented by solid lines with the 95% confidence limits for the individuals indicated
about them. The reduced major axis (RMA) regression lines for the comparative samples are the dashed lines. Homo sapiens RMA
formula: y p 4.833x 3.279; Pan RMA formula: y p 5.429x 33.197.

variable because of its behavioral, biological, and ecological
correlates (Calder 1984; Damuth and MacFadden 1990; Jung-
ers 1985; Millien and Bovy 2010; Schmidt-Nielsen 1984).
Across taxa, body mass is generally considered to be the most
appropriate measure of an animals overall size (Darveau et
al. 2002; Garcia and da Silva 2006; West, Brown, and Enquist
1997), but predicting body mass from skeletons involves the
introduction of a level of error that some find problematic
(Smith 1996). It has been shown that among primates, limb
articular dimensions, especially articular breadths, are among
the best predictors of body mass (Jungers 1990; McHenry
1992; Ruff 2003; Ruff, Walker, and Teaford 1989). In esti-
mating body mass for A.L. 288-1 using an all-hominoid re-
gression, Jungers (1990) found that femoral head diameter
showed the lowest standard error (SE) of the estimate and
lowest percentage predicted error of six lower limb articular
measurements, and importantly, McHenry (1992) found that
human regression formulas likely outperformed cross hom-
inoid predictive formulas in estimating fossil hominin body
mass. For this reason, a recent human least squares regression
formula (arguably the most appropriate formula for predic-
tion; see Hens, Konigsberg, and Jungers 2000; Jungers 1982;
Martin, Genoud, and Hemelrijk 2005; Smith 2009) from Mc-
Henry (1992) is used here to predict body mass from femoral
head diameter.
From this, the mean body mass of Australopithecus afarensis
is estimated at ca. 41 kg, while those of Australopithecus af-
ricanus, Australopithecus (P.) robustus, and Australopithecus
(P.) boisei are slightly smaller, at ca. 37.3, 37.0, and 38.5 kg,
respectively. In contrast, early (ca. 1.81.5 mya) Homo is
found to be considerably (ca. 33%) heavier, at ca. 54.5 kg.
This is close to the mean of the recent low-latitude human
sample (55.1 kg) although less heavy than that of recent high-
latitude humans (59.9 kg) and considerably lighter than Homo
neanderthalensis (ca. 73.6 kg) or late Pleistocene Homo sapiens
(64.1 kg). These and other differences are best visualized in
the box plots in figure 6. There is a clear dichotomy in es-
timated body mass between the members of the genera Aus-
tralopithecus and Paranthropus (on the left side of the graph)
and Homo on the right, a result similar to that obtained by
Pontzer (2012). Note, however, that while the earliest undis-
puted members of the genus Homo (at least those that pre-

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Holliday Body Size and Shape in Early Homo
Figure 6. Box plots of estimated body mass for fossil hominin and recent human samples.
serve a femoral head) are larger than almost all of the indi-
vidual australopiths, overall size in early Homo remains
smaller than that of Holocene or late Pleistocene H. sapiens
with the exception of modern-day Pygmoid groups, who
themselves show a reduction in body size likely due to selec-
tion related to climate, diet, high mortality rates, and/or island
biogeography (Bailey 1991; Cavalli-Sforza 1986; Hiernaux and
Froment 1976; Kuzawa and Bragg 2012; Lomolino 2005; Mig-
liano and Guillon 2012; Migliano, Vinicius, and Mirazon Lahr
2007).
Because of the importance of body mass in biology, there
is a rich literature on the estimation of body mass in fossil
hominins (Hartwig-Scherer 1993; Jungers 1990; Kappelman
1996; McHenry 1992; Ruff 1990; Ruff, Trinkaus, and Holliday
1997). As mentioned before, some (e.g., Smith 1996) feel that
because of the introduction of prediction error, body mass
in fossils should not be estimated but rather comparable mea-
surements that are known to be highly correlated with body
mass should be compared (although this alternative method
is not without its own problems; see Smith 1980). It should
therefore be noted that while body mass estimation formulas
based on femoral head size show low SEs of the estimate,
they are subject to error and interpretation.
One potential problem with body mass estimation from
femoral head size lies in the apparent difference in relative
femoral head size between Australopithecus (smaller femoral
heads) and Homo (larger femoral heads). Given this differ-
ence, should body mass in Australopithecus be estimated from
the femoral head using a nonhuman-hominoid formula or
rather a human and/or an all-hominoid one? In the smaller-
body size range, this question is largely academic, but it be-
comes problematic for larger-sized australopiths. For exam-
ple, two estimates of body mass from femoral head diameter
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S339
for the small A.L. 288-1 specimen, based on the all-hominoid
versus human regressions, are both 27.9 kg according to Mc-
Henry (1992). Likewise, Jungers (1990) arrives at a similar
estimate of 29.6 kg (only 6% higher) using a nonhuman-
hominoid formula. In contrast, McHenrys (1992) least
squares femoral head estimate of body mass for the larger
A.L. 333-3 specimen is 50 kg (based on a human formula),
and 37% higher, at 68.6 kg, for an all-hominoid formula.
Worse still, the nonhuman-hominoid least squares formula
reported in Jungers (1991b) produces an estimated body mass
of 81.9 kg for the specimen, 64% higher than McHenrys
(1992) human estimate. The body mass estimates reported
here must therefore be approached with caution. Despite this
caveat, other researchers, using different means of body mass
estimation, have found a similar dichotomy in body mass
between australopiths and Homo erectus (Kappelman 1996;
McHenry 1988; and see Pontzer 2012).
For recent humans, in addition to body mass, stature (or
standing height) is also frequently used as a proxy for body
size (Auerbach and Sylvester 2011; Malina et al. 2004; Ruff
2007; Zakrewski 2003). The problem with stature as a mea-
surement is that with skeletal data it is always estimated with
error. Given potential differences in limb : trunk proportions
between living humans and at least some australopiths (Fel-
desman, Kleckner, and Lundy 1990; Franciscus and Holliday
1992; Jungers and Stern 1983; but see Wolpoff 1983), and
given the fact that cranial height for most fossil hominins is
shorter than that of modern humans (Graves et al. 2010; Ruff
and Walker 1993), it is probably better to avoid estimating
stature in fossil hominins and instead analyze one of its major
constituents. In this study, femoral length is used as a proxy
for stature. While an imperfect reflection of stature, it is the
longest limb bone making up a portion of stature, and among
 S340
the long bones, it tends to show the lowest SE of the estimate
in stature prediction (Trotter 1970).
The distribution of femoral lengths of fossil and recent
hominins is shown in the box plot presented in figure 7. As
was the case with body mass, there is a dichotomy between
Australopithecus on the one hand (here represented solely by
the species A. afarensis) and Homo on the other. All of the
undisputed early Homo specimens fall well within the size
range of recent humans, and most have femora that are longer
than those of modern-day Pygmoid peoples (the early
Homo mean and median lie just below a single large Pyg-
moid outlier). In contrast, two of the five A. afarensis spec-
imens fall well below the modern human range of variation,
and for the three who fall within the modern human range
of variation, none falls outside the Pygmoid range of var-
iation.
Summary
With regard to the emergence of our genus, the postcranial
morphology of early Homo appears to have been mosaic in
nature. Of the features examined here, the most significant
difference between Australopithecus and early Homo is an in-
crease in body size as reflected in both body mass and stature
(importantly, this finding excludes OH 62, the femoral length
of which remains unknown). In contrast, differences in limb
proportions and body shape between the two genera are more
nuanced. For example, the differences in lower limb length
relative to overall size between Australopithecus and early
Homo are not as great as was once thought but rather the
relatively short femora of some specimens of both genera (e.g.,
Figure 7. Box plots of femoral length for fossil hominin and recent human samples.
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Current Anthropology Volume 53, Supplement 6, December 2012
A.L. 288-1 or LB1) appear to be primarily a function of their
diminutive body size (Franciscus and Holliday 1992; Holliday
and Franciscus 2009; Pontzer 2012). Likewise, while the
humero-femoral indices of early Homo are not as elevated as
those observed in Australopithecus afarensis (or likely those
of Australopithecus africanus; see McHenry and Berger 1998a,
1998b), it is evident that at least some specimens referred to
early Homo (e.g., Dmanisi 4167/4507) have longer humeri
relative to their femora than do later members of the genus.
This suggests that contra Asfaw et al. (1999), modern hu-
manlike humero-femoral indices were likely not characteristic
of hominins 2.5 mya but rather the low index values calculated
for the taxonomically unassigned BOU-VP-12/1 specimen are
the result of erroneous reconstruction of at least one of the
specimens limb bone lengths (most likely its humerus).
It is more difficult to evaluate relative forearm length in
Homo given a dearth of individuals who preserve antebrachial
elements. In general the genus Australopithecus, including the
late species Australopithecus sediba (Berger et al. 2010), ap-
pears to have had relatively longer forearms than modern
humans. At the same time, however, these forearms were
considerably shorter than those of Pan. While one must re-
main mindful that KNM-WT 15000 is a juvenile, its predicted
radius length relative to the length of its femur falls near the
upper margins of the combined late Pleistocene/Holocene
Homo sapiens sample, which may suggest that early members
of the genus Homo, including Homo ergaster/Homo erectus,
were characterized by longer forearms than later members of
the genus.
In terms of relative pelvic breadth, early Homo is similar
to Australopithecus in that it is characterized by wide bi-iliac
Holliday Body Size and Shape in Early Homo
breadths. This plesiomorphic character is also observed in
middle Pleistocene Homo and Homo neanderthalensis, sug-
gesting that the relatively narrow bi-iliac breadth seen in peo-
ple today is an autapomorphy of H. sapiens.
What emerges from these analyses, then, is that the origin
of the genus Homo is not a marked postcranial shift from an
australopith-like morphological pattern to an H. sapienslike
one, but as has been noted by others (e.g., Berger et al. 2010;
Kivell et al. 2011; Lordkipanidze et al. 2007; Pontzer 2012;
Pontzer et al. 2010), it is more complex. For example, lower
limb length appears to be as long in Australopithecus as it is
in early Homo once one corrects for differences in body size.
This brings up the fact that one long-recognized difference
between the australopiths and Homo that remains borne out
by the current analyses is that Homo tends to be much larger
in body size than Australopithecus or Paranthropus. This pat-
tern holds despite recent work (Graves et al. 2010; Ohman
et al. 2002) finding that the original projection of the adult
stature of KNM-WT 15000 (Ruff and Walker 1993) was far
too tall, or the revelation that many of the 1.77-million-year-
old Homo specimens from Dmanisi were of relatively dimin-
utive size (Lordkipanidze et al. 2007), or even the discovery
of relatively long-legged australopiths such as KSD-VP-1/1
(Haile-Selassie et al. 2010). It is a robust pattern: while on
average smaller than many people today, early Homo was
significantly larger than the australopiths in both body mass
and stature. And because among mammals, home-range size
is to a great extent a function of body size (Eisenberg 1990;
Harestad and Bunnell 1979; Kelt and Van Vuren 2001; McNab
1963; Vieira and Cunha 2008), it is almost certain that Homo
ranged more widely than Australopithecus, a notion under-
scored by the presence of Homo (and apparent lack of Aus-
tralopithecus or Paranthropus) outside of Africa.
Increased ranging in early Homo is most likely tied to an
increase in carnivory (Aiello and Key 2002; Martin 1981;
McHenry 1994; OConnell, Hawkes, and Blurton Jones 1999).
First, mammalian carnivores need larger home ranges than
comparably sized herbivores or omnivores (Gittleman and
Harvey 1982; Harestad and Bunnell 1979; Lindstedt, Miller,
and Buskirk 1986) and tend to exploit much greater day
ranges than noncarnivores as well (Carbone et al. 2005). Sec-
ond, larger bodies (and bigger brains; Aiello and Wheeler
1995) would likely only be made possible via an increase in
nutritional quality of the diet, and animal muscle and fat are
very high-caloric food sources known to have been exploited
by early Homo. Evidence in favor of increased carnivory in
early Homo include the 1.7-million-year-old partial H. erg-
aster/H. erectus skeleton KNM-ER 1808, argued by Walker,
Zimmerman, and Leakey (1982) to show pathological bone
suggestive of hypervitaminosis A, which can be caused via
ingestion of carnivore livers. Likewise, from an archaeological
perspective, the Acheulean Industry, which is now known to
date as early as 1.76 mya (Lepre et al. 2011), is temporally
correlated with zooarchaeological evidence from multiple sites
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S341
indicative of a shift toward consistent early access to carcasses
by hominins (Cachel and Harris 1998).
Despite the important adaptive implications surrounding
increased ranging activity in early Homo, aside from a few
anatomical features of the foot (DeSilva et al. 2012; Pontzer
et al. 2010; Zipfel et al. 2011), pelvis (Berger et al. 2010; Kibii
et al. 2011), and some features related to head carriage (Bram-
ble and Lieberman 2004), there is little evidence of a major
locomotor shift between Australopithecus and Homo. Instead
it appears that Homo augmented an existing australopith pat-
tern almost exclusively via an increase in body size (and see
Pontzer 2012).
Following these observations, at least from a postcranial
adaptive-zone perspective, there are insufficient data to war-
rant the removal of Homo habilis and Homo rudolfensis from
the genus Homo as recommended by Wood and Collard (Col-
lard and Wood 2007; Wood and Collard 1999a, 1999b). Of
the cranial remains assigned to these two species, only two
(OH 62 and KNM-ER 3735) have associated postcrania, and
in both cases, the postcranial remains are extremely frag-
mentary and therefore relatively uninformative (Haeusler and
McHenry 2007). In this light, if any of the long (but isolated)
femora dating to ca. 1.8 mya in Africa is referred to either of
these species, then we have probably underestimated both
body size and lower limb length in these two taxa. Addition-
ally, because H. ergaster/H. erectus may have retained a some-
what elongated upper limb and an equally broad pelvis, it
appears that many of the perceived proportional differences
between H. habilis and/or H. rudolfensis on the one hand and
specimens assigned to H. ergaster or H. erectus on the other
are diminished.2
As for the monophyletic nature of the genus Homo, while
some recent cladistic analyses (e.g., Cameron and Groves
2004) fail to cluster H. habilis and H. rudolfensis with H.
sapiens to the exclusion of any australopith taxon, it is im-
portant to note that Cameron and Grovess (2004) result is
entirely due to the inclusion of Kenyanthropus platyops in the
analyses, a paleospecies whose taxonomic validity has been
questioned (White 2003). Simply removing this species from
Cameron and Grovess (2004) analysis makes Homo (sensu
lato) a monophylum. More importantly, the most rigorous
cladistic analyses of fossil hominins done to date, including
some analyses that include K. platyops (Strait and Grine 2004)
show Homo (sensu lato) to be a monophylum. In any case,
the close phylogenetic relationship of Homo and the austral-
opiths (it seems unlikely that they split much before 2.5 mya)
probably lent itself to some (albeit low) level of interbreeding
between these taxa, which in turn could produce confusing
patterns of synapomorphy (Holliday 2003, 2006; Jolly 2001).
In conclusion, then, while the emergence of Homo (sensu
2. There are also numerous craniodental characters that support the
inclusion of Homo habilis and Homo rudolfensis in the genus Homo
(Anton 2012; Bromage, Schrenk, and Zonneveld 1995; Kimbel, Johanson,
and Rak 1997; Rightmire and Lordkipanidze 2009).
 S342
lato) is an important chapter in human evolutionone likely
involving increased ranging, carnivory, and encephalization
it is also the case that we now have the resolution in the fossil
record to actually capture the gradual mosaic nature of that
emergence. The long-standing idea of the origin of Homo
(sensu stricto) as a punctuated event involving the geologically
rapid emergence of hominins who (at least from the neck
down) were H. sapienslike seems less applicable now than
it did a decade ago.
Acknowledgments
I would like to thank Leslie Aiello and Susan Anton for in-
viting me to participate in the Wenner-Gren symposium, and
to them and all the other participants, my thanks for making
the symposium such a phenomenal and unforgettable expe-
rience. Thanks also to Tim Weaver for supplying a bi-iliac
breadth estimate for Tabun C1 and to Erik Trinkaus for insight
into the morphology of the La Chapelle-aux-Saints pelvis.
Thanks, too, to Campbell Rolian for generously providing
some of the Pan and Gorilla data included in these analyses.
This work was supported in part by the National Science
Foundation, the Leakey Foundation, Tulane University, and
the University of the Witwatersrand.
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