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Journal of Biogeography (J. Biogeogr.) (2009)
SPECIAL
PAPER
Evolution of the second orangutan:
phylogeny and biogeography of hominid
origins
John R. Grehan
1
* and Jeffrey H. Schwartz
2
1
Buffalo Museum of Science, Buffalo, NY and
2
Departments of Anthropology, and History
and Philosophy of Science, University of
Pittsburgh, Pittsburgh, PA, USA
ABSTRACT
Aim
To resolve the phylogeny of humans and their fossil relatives (collectively,
hominids), orangutans (
Pongo
) and various Miocene great apes and to present
a biogeographical model for their differentiation in space and time.
Location
Africa, northern Mediterranean, Asia.
Methods
Maximum parsimony analysis was used to assess phylogenetic
relationships among living large-bodied hominoids (= humans, chimpanzees,
bonobos, gorillas, orangutans), and various related African, Asian and European
ape fossils. Biogeographical characteristics were analysed for vicariant
replacement, main massings and nodes. A geomorphological correlation was
identified for a clade we refer to as the ‘dental hominoids’, and this correlation
was used to reconstruct their historical geography.
Results
Our analyses support the following hypotheses: (1) the living large-
bodied hominoids represent a monophyletic group comprising two sister
clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas
(collectively, the African apes); and (2) the human–orangutan clade (dental
hominoids) includes fossil hominids (
Homo
, australopiths,
Orrorin
) and the
Miocene-age apes
Hispanopithecus
,
Ouranopithecus
,
Ankarapithecus
,
Sivapithecus
,
Lufengpithecus
,
Khoratpithecus
and
Gigantopithecus
(also Plio-Pleistocene of
eastern Asia). We also demonstrate that the distributions of living and fossil
genera are largely vicariant, with nodes of geographical overlap or proximity
between
Gigantopithecus
and
Sivapithecus
in Central Asia, and between
Pongo
,
Gigantopithecus
,
Lufengpithecus
and
Khoratpithecus
in East Asia. The main
massing is represented by five genera and eight species in East Asia. The dental
hominoid track is spatially correlated with the East African Rift System (EARS)
and the Tethys Orogenic Collage (TOC).
Main conclusions
Humans and orangutans share a common ancestor that
excludes the extant African apes. Molecular analyses are compromised by
phenetic procedures such as alignment and are probably based on primitive
retentions. We infer that the human–orangutan common ancestor had
established a widespread distribution by at least 13 Ma. Vicariant
differentiation resulted in the ancestors of hominids in East Africa and various
primarily Miocene apes distributed between Spain and Southeast Asia (and
possibly also parts of East Africa). The geographical disjunction between early
hominids and Asian
Pongo
is attributed to local extinctions between Europe and
Central Asia. The EARS and TOC correlations suggest that these
geomorphological features mediated establishment of the ancestral range.
*Correspondence: John R. Grehan, Buffalo
Museum of Science, 1020 Humboldt
Parkway, Buffalo, NY 14211-1293, USA.
E-mail: jgrehan@sciencebuff.org
Keywords
Hominid, hominoid, human origin, orangutan, panbiogeography,
Sivapithecus
,
systematics, Tethys, vicariance.
ª
2009 Blackwell Publishing Ltd
www.blackwellpublishing.com/jbi
1
doi:10.1111/j.1365-2699.2009.02141.x
J. R. Grehan and J. H. Schwartz
INTRODUCTION
molecularly based theory of the relationship among large-
bodied hominoids and the biogeography of these primates
(Schwartz, 1987, 2005; Grehan, 2006b).
Biogeographical reconstructions of a common African
origin of hominids and African apes have been dominated
by Darwin’s (1859) assumption that vicariant (spatially
disjunct) fossil localities are historically connected by a series
of discrete migrations from common centres of origin, as well
as by Matthew’s (1915) assumption that centres of origin and
migration can be read – literally – from the fossil record
(Heads, 2005). Localities with older fossils or basal lineages are
generally assumed to represent actual sites of earlier occur-
rence, and multiple area relationships are attributed to
multiple migrations or reciprocal migrations back and forth
according to theoretical parsimony criteria (e.g. Beard, 2006;
Fleagle & Gilbert, 2006; Heesy
et al.
, 2006; Folinsbee & Brooks,
2007).
The fossil record of modern African apes is limited to a few
putative chimpanzee teeth dated to
c
. 0.5 Ma (McBrearty &
Jablonski, 2005). In the absence of older fossils either of any
African ape or of a presumed common African ape–hominid
ancestor, hominoid systematists have attempted to link the
origin of early fossil hominids in Africa with various Miocene
fossil apes in Eurasia through a series of hypothetical dispersals
that Begun (2001) characterized as ‘very complicated’. Begun
(2001) himself theorized an initial dispersal of hominoids into
Europe where they diverged to give rise to the Asian apes as
well as to a hypothetical common ancestor of humans and
African apes that migrated back into Africa
c
. 9 Ma. Several
similar biogeographical scenarios have been proposed accord-
ing to differing parsimony hypotheses about the historical
interrelationships between centres of origin and dispersal,
phylogeny and area (Moy
`
Sol
`
&K¨hler, 1993; Begun
et al.
,
1997, 2003; Begun & G¨ le¸ , 1998; Miyamoto & Young, 1998;
Stewart & Disotell, 1998, 1999; Moy
`
Sol
`
&K¨hler M., 1999;
Begun & Nargolwalla, 2004; Cote, 2004; Begun, 2005; Harri-
son, 2005; Pickford & Senut, 2005; Pickford, 2006; Folinsbee &
Brooks, 2007). Recently discovered fossils have been inter-
preted as providing evidence of African ape-like hominoids in
Africa
c
. 12.5–10 Ma (Pickford & Senut, 2005; Suwa
et al.
,
2007) but their lack of African ape synapomorphies brings this
interpretation into serious question.
These complicated and convoluted dispersal models are not
grounded in empirical evidence, but rather on preconceived
notions about the existence and locations of centres of origin.
Following Darwin (1859), the underlying assumption is that an
ancestor’s distribution is geographically narrow with respect to
the distributions of presumed descendants located in different
(vicariant) geographical areas. The problem with this precon-
ception, however, is that the very ability of individual
descendants to move between different locations obviates the
very criterion – geographical isolation – that is also invoked as
essential for their differentiation and speciation. An alternative
model of vicariant differentiation (Croizat, 1958) suggests that
ancestral dispersal occurs before the differentiation of descen-
dant taxa, with the ancestor establishing a widespread
The order Primates subsumes three major groups: prosimians
(broadly, lemurs, lorises and bushbabies), anthropoids (mon-
keys, apes and humans) and tarsiers, with conflicting evidence
for the grouping of the latter with prosimians rather than
anthropoids (Schwartz, 1986; Groves, 1991, 2006). Anthro-
poids are traditionally subdivided into Platyrrhini (New World
monkeys) and Catarrhini (Old World monkeys, apes and
humans). The catarrhine superfamily Hominoidea includes the
large-bodied hominoids or great apes (hominids, orangutans
and African apes) as well as the small-bodied hominoids or
lesser apes (Hylobatidae, i.e. gibbons and siamangs; Schwartz,
1986). For over a century the great apes, which were thought to
be closely related, were allocated to the family Pongidae, but
the last three decades have witnessed a shift in favour of
interpreting overall molecular similarity as indicating that the
African apes (chimpanzees, bonobos and gorillas) and now
chimpanzees and bonobos alone are most closely related to
humans (Marks, 2003; Schwartz, 2007; Schwartz & Maresca,
2007; Senut, 2007). The latter presumption has led some
biologists to refer to humans as a third chimpanzee (Diamond,
1993) and others to place chimpanzees and all hominids in the
same genus,
Homo
(Goodman
et al.
, 1998).
Most palaeoanthropologists taxonomically accommodate a
human–African ape relationship either by restricting Pongidae
to the orangutan and its fossil relatives while placing African
ape and hominid genera within Hominidae, or by including all
large-bodied hominoids in Hominidae (Andrews & Bernor,
1999) within which humans and their fossil relatives are placed
in either the subfamily Homininae or the tribe Hominini (e.g.
Goodman, 1996; Cameron, 1997; Begun & G¨ le¸ , 1998; Leakey
et al.
, 2001; Begun, 2004; Pilbeam & Young, 2004; Andrews &
Harrison, 2005; Folinsbee & Brooks, 2007). Here we follow an
alternative protocol that restricts the family Hominidae to
include only humans and their fossil relatives to the exclusion
of their closest living great ape relative(s) (cf. Schwartz, 1986,
2007; Hill & Ward, 1988; Strait & Grine, 2004; Tuttle, 2006;
Strait
et al.
, 2007).
Pilbeam & Young’s (2004) assertions notwithstanding, a
presumed human–African ape relationship is phylogenetically
quite problematic. Neither of two oft-cited morphological
studies claiming to corroborate the interpretation of molecular
data as supporting a close relationship between chimpanzees
and humans (Shoshani
et al.
, 1996; Begun
et al.
, 1997) took
into consideration or provided justification for excluding most
of the morphological features that have been documented as
being shared uniquely by humans and orangutans (see
discussions in Schwartz, 2005; Grehan, 2006a). Of further
note, the studies by Begun
et al.
(1997) and Cameron (1997)
claiming support for a human–chimpanzee relationship did
not include any extinct species of
Homo
. Clearly, however,
given the considerable morphological evidence in support of it,
the hypothesis of a closer relationship between humans and
orangutans than between humans and either extant African
ape genus has major implications for evaluating both the
2
Journal of Biogeography
ª
2009 Blackwell Publishing Ltd
Phylogeny and biogeography of hominid origins
geographical distribution that encompasses the combined
ranges of all vicariant descendants (Craw
et al.
, 1999).
We take the position that integrating theories of relatedness
between fossil and living taxa with those concerning biogeog-
raphy can be accomplished only if shared–derived morpho-
logical similarities are first delineated and then used to
generate and subsequently test theories of relatedness (Sch-
wartz, 1987, 2005). Here we present a morphological analysis
of living and various fossil large-bodied hominoids in order to
test alternative theories of relationship among and between
them. We then integrate the resulting hypothesis of relatedness
into a historical biogeographical model that considers the
extent of vicariant replacement among member taxa as a
predictor of the geographical range of the most recent
common ancestor. We also examine possible tectonic corre-
lations with the ancestral hominid range as a biogeographical
calibration of the minimal age of origin and dispersal of the
ancestor of a hypothesized common ancestor of hominids,
orangutans and their closest fossil ape relatives.
Characters and phylogenetic analysis
Our analysis of potential phylogenetic relationships among
extant large-bodied hominoids is based on a character matrix
(see Appendix S1 in Supporting Information) comprising
structural, behavioural and physiological features (Schwartz,
1987, 1988, 2004a, 2005, 2007; Grehan, 2006a). In contrast to
more commonly published approaches, e.g. Strait & Grine
(2004) in which purported cladistic analysis often claims
synapomorphy within the ingroup in spite of the fact that the
feature is also common in the outgroup, the vast majority of
the features we propose as potential synapomorphies between
humans and orangutans are not represented in any outgroup
species. Only for enamel thickness (character 59 for living taxa,
character 1 for all taxa) was a feature also present in the
outgroup considered sufficiently rare as to represent an
independent origin (see Appendices S1 & S2).
As much as possible, and in contrast to the practice in
primate molecular studies (e.g. Ruvolo, 1997) and increasingly
so in morphological studies (e.g. Begun & G
¨
le¸ , 1998;
Lockwood
et al.
, 2004; Strait & Grine, 2004), we included
whenever possible a significant number if not all taxa in the
outgroup, rather than only a few selected taxa. Where
character states involved quantitative differences (such as in
relative or absolute size and volume or angle and orientation),
we limited the informative state to the two most derived
conditions shared by any two taxa. The values for the other
taxa are documented for each character in Appendices S1 and
S2. For some characters we accepted a threshold value
as proposed in other studies in support of derived states
shared between humans and chimpanzees or humans and
African apes.
Our analysis of relationships between living and fossil taxa
is based on a character matrix limited to hard-tissue
characters that have been sufficiently well described in the
literature to permit verification, and whose claimed charac-
ter states as well as unique occurrence within a large-bodied
hominoid clade we could corroborate via a broad outgroup
comparison (see Appendix S2). Our treatment of some
hominid taxa with multiple species (e.g.
Australopithecus
)as
single taxonomic units was sufficient to address their
relationship to extant great apes while not precluding the
possibility that they may be paraphyletic, or even polyphy-
letic, with respect to
Homo
. For the purposes of our analysis
we accept the following hypotheses: Anthropoidea (New and
Old World ‘higher’ primates) constitutes a monophyletic
group that subsumes the monophyletic groups Platyrrhini
(New World monkeys) and Catarrhini (Old World
monkeys and hominoids), and that among the hominoids
the monophyletic Hylobatidae (gibbons and siamangs) is the
sister group of large-bodied hominoids. These hypotheses
have withstood continual testing and are highly corroborated
(e.g. Delson & Andrews, 1975; Groves, 1986; Schwartz,
1986; Shoshani
et al.
, 1996; Schwartz & Yamada, 1998),
although relationships of taxa within any clade may be
contested.
MATERIALS AND METHODS
Taxonomic groups
Accepted hominids are represented here by
Homo
(Schwartz &
Tattersall, 2001, 2003, 2005) and the australopiths (
Australop-
ithecus
,
Paranthropus
; Schwartz & Tattersall, 2005). We also
include in our analysis four other proposed hominids:
Orrorin
(Senut
et al.
, 2001),
Kenyanthropus
(Leakey
et al.
, 2001),
Ardipithecus
(White
et al.
, 1994, 1995) and
Sahelanthropus
(Brunet
et al.
, 2002). We do so primarily because of the
publicity they have received and the claims made for their
hominid status. But for the latter three, we do so with the
caveat that only when their discoverers make at least the type
specimens available for scrutiny by others can independent
verification of the published descriptions and interpretations
be possible (see commentaries by Schwartz, 2004b; Schwartz &
Tattersall, 2005).
Our sample of fossil apes includes only those taxa for
which substantial morphological evidence has already been
provided in support of their cladistic membership within a
large-bodied hominoid clade:
Hispanopithecus
,
Ouranopithe-
cus
,
Ankarapithecus
,
Sivapithecus
,
Gigantopithecus
,
Lufengpi-
thecus
,
Khoratpithecus
and
Dryopithecus
(e.g. Schwartz, 1990,
1997; Moy` Sol` &K¨hler, 1993; Stewart & Disotell, 1998;
Begun, 2005). Other extinct large-bodied hominoids are
either apparently more basal to these groups or insufficiently
known skeletally to adequately resolve their phyletic position
(see Begun, 2001). The fossil taxa
Langsonia
from Vietnam
(Schwartz
et al.
, 1995) and ‘
Sivapithecus
’ from Nepal (Mun-
the
et al.
, 1983) are not included here because they lack
diagnostic features that can potentially suggest relationships
beyond their membership within a hypothesized clade of
dentally thick-enamelled apes. Extant ingroup taxa in our
analysis are the species
Homo sapiens
and the great ape genera
Gorilla
,
Pan
, and
Pongo
.
Journal of Biogeography
3
ª
2009 Blackwell Publishing Ltd
J. R. Grehan and J. H. Schwartz
Characters used in our analyses emerged from a collation
that initially consisted of hundreds of features generated for,
and used in, various studies of human–great ape relationships.
Most of these features, however, were found to be irrelevant to
the question of relationships among the large-bodied homi-
noids (e.g. they appeared to be plesiomorphic), incorrectly
identified or unverifiable (Schwartz, 1987, 1988, 2005; Grehan,
2006a; see Appendices S3–S5), even after requests for clarifi-
cation addressed to the original authors. Elsewhere we have
proposed up to 42 characters as uniquely shared between
humans and orangutans, but of these only 28 are included here
as particularly well corroborated at this time (see Appen-
dix S1). This total does not include an additional seven
characters proposed by other researchers for humans and
orangutans that require further exploration (Grehan, 2006a).
Of the 28 human–orangutan characters supported here, 15
have been accepted by various authors who embrace a human–
chimpanzee relationship (Andrews, 1987; Groves, 1986, 1987;
personal communication; Graham, 1988; Thiranagama
et al.
,
1991; Shoshani
et al.
, 1996; Strait & Grine, 2004). The other 13
characters have essentially been dismissed out of hand or
ignored without proper evaluation by others (Grehan, 2006a).
We acknowledge that some primate biologists and systematists
object to using characters with a presumed functional role
because they may be the result of selection independent of
phylogeny. In the absence of empirical evidence, this objection
is rhetorical. We take the view that such assumptions are not
relevant to phylogenetic analysis [because they incorrectly
embody use–disuse arguments in which the assumption ‘form
follows function’ predominates (Croizat, 1964; Schwartz,
2005)] and that the most highly corroborated hierarchically
nested set of derived characters yields the most probable
phylogenetic relationship (Nelson & Platnick, 1981).
In order to maintain analytical comparability between our
analyses and other studies on large-bodied hominoid relation-
ships, we used maximum parsimony analysis with paup*
4.0b10 (Swofford, 2005) and tnt (sponsored by the Willi
Hennig Society; Goloboff
et al.
, 2008) to identify diagnostic
synapomorphies. We included bootstrap (50% majority rule)
and Bremer support analysis as two widely used measures of
tree viability. An exhaustive search was made for extant taxa
(humans and the great apes) in Analysis A, and between extant
taxa and unproblematic fossil hominids (australopiths and
Homo
) in Analysis B. A heuristic search was made in Analysis
C for all extant and fossil taxa considered in this study. In each
analysis, characters exclusive to small- and large-bodied
hominoids were included to illustrate their monophyly in
relation to monkeys. The same analysis was also carried out in
Analysis D where only those fossil taxa with sufficient
informative characters to provide resolution of relationships
within the ‘dental hominoid clade’ (defined in Results) were
included. This technique recognizes that missing data may
increase the number of equally parsimonious trees as well as
result in the production of spurious cladograms (Ebach &
Ahyong, 2001). Unresolved relationships in Analyses C and D
were identified by strict consensus.
Biogeographical analysis
The minimum-spanning tree (track) method (Craw
et al.
,
1999) was used to reconstruct the spatial connection between
the disjunct and vicariant distributions of hominids and non-
hominid members of the dental hominoid clade. Disjunct
localities of each taxon, whether living or fossil, were linked
together as a minimal spanning tree, and these tracks were then
connected to each other by additional minimal spanning links
between nearest localities. The spatial structure of the track was
characterized with respect to the vicariant replacement of taxa,
the intersection of two or more individual tracks (nodes) and
main massings (geographical concentrations of diversity
whether genetic, morphological, taxonomic, etc.). These spatial
features are used to provide an evolutionary model for the
differentiation of hominids and their nearest living and fossil
relatives.
Spatial overlap (geological correlation) with tectonic or
geomorphological features was examined in order to gener-
ate a historical model for estimating the minimum diver-
gence age and distribution range of the last common
ancestor. The distribution beyond Africa of species of
Homo
is generally considered to have resulted from one or more
range expansions following an African origin of the genus.
Since
Homo
is widespread and sympatric with respect to
all other members of the dental hominoid clade, the
biogeography of
Homo
lies outside the scope of the present
analysis.
Taxa
(1) Ankarapithecus
Known from a single species, formerly known as
Sivapithecus
meteai
Ozansoy, 1957, from 10.7 to 10.6 Ma in the Sinap
Formation north of Yassi¨ren in central Turkey (Fig. 1a;
Andrews & Tekkaya, 1980; Alpagut
et al.
, 1996; Lunkka
et al.
,
1999). The specimens comprise a mandible and skull frag-
ments that present a tall, wide and anteriorly facing zygoma
(cheekbone), marked alveolar prognathism combined with a
short upper face, tall, ovoid, superiorly rimmed but not widely
separated orbits, and a long, slit-like incisive foramen situated
anteriorly in the palate – all of which suggest affinity with the
orangutan (Andrews & Cronin, 1982).
(2) Ardipithecus
Two species from the Middle Awash region of Ethiopia
(Fig. 1a). The holotype of
Ardipithecus ramidus
(White
et al.
,
1994) comprises a set of ‘associated teeth from one individ-
ual’ and the paratype series has various associated cranial
fragments, two partial cranial bases, a juvenile mandible and
associated and isolated teeth dated
c
. 4.4 Ma. Molars are
characterized as having absolutely and relatively thinner
enamel than those in
Australopithecus
(White
et al.
, 1994,
1995). The holotype of
Ardipithecus kadabba
Haile-Selassie,
4
Journal of Biogeography
ª
2009 Blackwell Publishing Ltd
Phylogeny and biogeography of hominid origins
Si
va
pithecus
derived features supporting their claim that
Australopithecus
bahrelghazali
Brunet
et al.
, 1995 from Chad is a hominid;
because independent study of the holotype was not permitted
(Schwartz, 2005), this taxon is not recognized here. Australo-
piths share many derived postcranial features with humans,
particularly with respect to bipedal locomotion (see review in
Schwartz, 2007), but many of their derived craniodental
features also characterize orangutans and their potential
extinct relatives (Schwartz, 2004a). If australopiths are more
closely related to humans than to any living great ape, the
features they share with the orangutan may represent primitive
retentions from the ancestor of a larger clade that subsumes
them all.
Ankarapithecus
L
u
fengpithecus
Sahelanthropus
Ardipithecus
Khoratpithecus
Kenyanthropus
Orrorin
(a)
(4) Dryopithecus
Dryop
it
hec
us
Three species from Europe (Fig. 1b)
c
. 9–12 Ma. Characterized
by the unique development of narrow, tall-crowned upper
central incisors and thin-enamelled molars with high dentine
penetrance.
Dryopithecus fontani
Lartet, 1856 from France and
Austria is represented by fragmentary mandibles (the holotype
is a mandible), isolated teeth and a humeral shaft (Begun,
1994).
Dryopithecus brancoi
Schlosser, 1901 from Rudab´nya,
Hungary is represented by the holotype (a left M
3
) plus
additional material, including a partial cranium (RUD 77),
tooth-bearing mandibles and maxillae and isolated teeth. The
holotype of
Dryopithecus crusafonti
Begun, 1992 from Can
Ponsic and El Firal, Spain comprises a poorly preserved left
maxilla and a separate but apparently associated left canine
fragment. In addition, 15 isolated teeth and a mandible have
recently been allocated to this taxon (Begun, 2002). Begun &
Kordos (1997) and Begun
et al.
(1997) have proposed
Dryopithecus
as the sister taxon of an australopith–African
ape clade.
Gigantopithecus
Pon
go
australopiths
(b)
Figure 1
Generalized distribution localities for fossil and living
hominoids (excluding
Homo
,
Pan
and
Gorilla
) included in this
study. (a) Fossil hominoids
Hispanopithecus
(Spain),
Ouranopi-
thecus
(Greece),
Ankarapithecus
(Turkey),
Sivapithecus
(Indo-
Pakistan),
Lufengpithecu
s (southern China),
Khoratpithecus
(Thailand),
Sahelanthropus
(Chad),
Ardipithecus
(Ethiopia),
Kenyanthropus
(Kenya) and
Orrorin
(Kenya). (b) Distribution of
extant (outline) and fossil
Pongo
in Southeast Asia, and fossil
hominoids
Dryopithecus
(Europe),
Gigantopithecus
(Indo-Pakistan
and eastern Asia) and the australopith hominids (East Africa).
(5) Gigantopithecus
A few incomplete lower jaws and numerous isolated teeth
allocated to three species:
Gigantopithecus blacki
von Koenigs-
wald, 1935 from south-eastern China and northern Southeast
Asia between 2.0 and 0.3 Ma (Pei & Woo, 1956; Simons &
Ettel, 1970; Ciochon
et al.
, 1990, 1996; Huang
et al.
, 1995;
Schwartz
et al.
, 1995; Zhao
et al.
, 2006),
Gigantopithecus
bilaspurensis
Simons & Chopra, 1969 (Simons & Chopra,
1968, 1969; Patnaik
et al.
, 2005) and
Gigantopithecus giganteus
(Simons & Chopra, 1968, 1969; Cameron, 1997, 2001, 2003)
from Indo-Pakistan (Fig. 1b)
c
. 7.5–7.8 Ma. The holotypes
comprise a right lower third molar for
G. blacki
, a right lower
second or third lower molar for
G. giganteus
(Kelley, 2002) and
a lower jaw lacking ascending rami for
G. bilaspurensis
(Simons & Chopra, 1968, 1969). The genus
Indopithecus
was
resurrected for the Indo-Pakistan species by Cameron (1997,
2001, 2003) because of perceived dental differences with
G. blacki
. Aside from a partial mandible for
G. blacki
and
G. bilaspurensis
,
Gigantopithecus
is known primarily from
isolated, often highly worn teeth.
Suwa & White, 2004 comprises part of a right mandible with
associated teeth, and various isolated teeth and postcranial
fragments, some from the same site as the holotype; dated to
c
. 5.2–5.8 Ma (Haile-Selassie, 2001; Haile-Selassie
et al.
,
2004).
(3) Australopithecus–Paranthropus
At least 10 species in East and South Africa (Fig. 1b)
c
. 4.5–
2.0 Ma (Schwartz & Tattersall, 2005). The relationships of
australopiths to one another are very uncertain (Schwartz,
2004a), and some australopiths appear to be more closely
related to
Homo
than others (Tattersall & Schwartz, 2000;
Strait & Grine, 2004). Brunet
et al.
(1995) did not delineate
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5
ª
2009 Blackwell Publishing Ltd
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Proctor, Human Recency and Race (2005).pdf
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