Paleobiology, 30(4), 2004, pp. 614–651
Reassessing hominoid phylogeny: evaluating congruence in the
morphological and temporal data
John A. Finarelli and William C. Clyde
Abstract.—The phylogenetic relationships of fossil and extant members of the primate superfamily
Hominoidea are reassessed by using both conventional (morphological) cladistic and stratocladistic (incorporating morphological and temporal data) techniques. The cladistic analysis recovers
four most parsimonious cladograms that distinguish postcranially primitive (‘‘archaic’’) and derived (‘‘modern’’) hominoid clades in the earliest Miocene of East Africa and supports distinct
hominine and pongine clades. However, the relationships among the pongines and hominine clades
and other Eurasian hominoids remain ambiguous and there is weak support (Bremer decay indices,
reduced consensus, and bootstrap proportions) for several other parts of the proposed phylogeny.
An examination of the partitioning of homoplasy across the two major hominoid clades recovered in the cladistic analysis indicates that the majority of the observed homoplasy resides in the
postcranially derived clade. An examination of the partitioning of homoplasy across anatomical
regions indicates that dental characters display a significantly higher level of homoplasy than postcranial characters. A rarefaction analysis demonstrates that the higher homoplasy associated with
the dental characters is not the result of sampling biases, indicating that postcranial skeletal characters are likely the more reliable phylogenetic indicators in the hominoids.
The branching order of the most parsimonious cladograms shows better than average congruence
with the observed ordering of first appearances in the fossil record, implying that the hominoid
fossil record is surprisingly good. As with morphologic parsimony debt, most of the stratigraphic
parsimony debt in these cladograms is associated with the ‘‘modern’’ hominoid clade. A stratocladistic analysis of the data recovers a single most parsimonious phylogenetic tree with a different
cladistic topology from the morphological cladogram. The most striking difference is the elimination of the postcranially primitive clade of hominoids in the early Miocene in favor of a pectinate
succession of taxa. The relative position of the late-appearing taxon Oreopithecus is also altered in
the stratocladistic hypothesis. Topological differences between the cladistic and stratocladistic hypotheses highlight two intervals of significant discord between the morphological and temporal
data—the early Miocene of eastern Africa and the late Miocene of Eurasia. The first discrepancy is
likely the result of poor preservation and morphological homoplasy in Morotopithecus, as the fossil
record in the early Miocene of eastern Africa for the ingroup is rather good. The second discrepancy
is likely the result of the unusual preservation conditions associated with the late Miocene hominoid Oreopithecus.
John A. Finarelli. Committee on Evolutionary Biology, University of Chicago, 1025 East Fifty-seventh
Street, Culver Hall 402, Chicago, Illinois 60637, and Department of Geology, The Field Museum of Natural
History, 1400 South Lake Shore Drive, Chicago, Illinois 60605. E-mail: [email protected]
William C. Clyde. Department of Earth Sciences, 56 College Road, University of New Hampshire, Durham,
New Hampshire 03824. E-mail: [email protected]
Accepted:
18 February 2004
Introduction
Despite a considerable research effort to refine our understanding of the evolutionary relationships among known fossil and extant
hominoid taxa, little consensus has emerged
(e.g., Begun 1992a,b, 1994, 1995; Begun et al.
1997a; de Bonis and Koufos 1993; Dean and
Delson 1992; Andrews et al. 1996; MacLatchy
et al. 2000). Although their use in phylogenetic
analysis remains controversial, temporal data
represent a potentially relevant class of information that is often overlooked (Fisher 1991,
1994; Huelsenbeck 1994; Wagner 1995). To
q 2004 The Paleontological Society. All rights reserved.
date no phylogenetic analysis of the Hominoidea has explicitly incorporated temporal
data in hypothesis testing, and no quantitative
evaluation of the congruence of the morphological and temporal data for the hominoid
fossil record has occurred. In an attempt to
clarify these evolutionary relationships and to
evaluate congruence across data types, a phylogenetic reassessment of the Hominoidea was
performed incorporating both morphological
and stratigraphic data.
Although cladistic hypotheses are evaluated with no explicit reference to time, all clad0094-8373/04/3004-0007/$1.00
DATA CONGRUENCE IN HOMINOID PHYLOGENY
ograms make implicit statements about the
relative time of divergence among taxa by the
order of branching events (Fisher 1991; Wagner 1995). The sequence of branching events in
a morphological cladistic hypothesis is often
harmonized with the fossil record of the ingroup through the creation of ‘‘ghost lineages,’’ artificial extensions of a taxon’s range beyond its observed first appearance in the fossil
record (Norell 1993). This approach essentially erases any discrepancy between the observed order of appearance events and the order implied by the hypothesis. Insofar as
ghost lineages explain away discrepancies between (stratigraphic) observation and (cladistic) hypothesis, they may be considered appeals to ad hoc support, analogous to the way
homoplasy is invoked to explain away morphological data that are incongruent with a
cladistic hypothesis (Fisher 1991, 1994).
Stratocladistics is a parsimony-based criterion that evaluates competing phylogenetic
hypotheses relative to both morphological
characters and a stratigraphic character derived from the stratigraphic record of the ingroup taxa (Fisher 1991, 1992, 1994). The morphological component is evaluated as in conventional cladistic analyses, where each instance of homoplasy imparts a unit of
‘‘morphological parsimony debt’’ upon the
hypothesis. In addition, each instance of incongruence between the stratigraphic data
and the hypothesis, where a taxon is predicted
to exist by the branching order yet is not observed, imparts a unit of ‘‘stratigraphic parsimony debt’’ (Fisher 1992). Stratocladistics
then sums the morphological and stratigraphic parsimony debt values of each hypothesis,
creating a single ‘‘total parsimony debt’’ value. The minimum total parsimony debt value
over the set of possible phylogenetic hypotheses determines the overall most parsimonious phylogenetic hypothesis (Fisher 1992,
1994).
Two recent studies have shown that stratocladistics may outperform conventional cladistic analyses in recovering evolutionary histories for fossil taxa. Working with published
cladistic analyses, Clyde and Fisher (1997)
noted significant increases in the fit of stratigraphic data to phylogenic hypotheses recov-
615
ered under stratocladistic analysis, without simultaneously observing significant decreases
in the fit of morphological data. In computer
simulations in which fossil records were created for hypothetical taxa with known evolutionary histories, stratocladistics significantly
outperformed conventional cladistic analysis
in recovering the true phylogenies (Fox et al.
1999). Additionally, in cases where neither
method was able to recover the correct phylogeny, the stratocladistic hypotheses more
closely matched the true phylogenies (Fox et
al. 1999).
Methods
Morphological Data
Thirteen fossil and five extant genera of the
primate superfamily Hominoidea constituted
the ingroup of this analysis. The morphological data matrix was modified from the character-by-taxon matrix of Begun et al. (1997a).
That study examined 240 morphological characters for eight fossil and five extant genera.
Modifications to the Begun et al. (1997a) data
set are discussed below. The character state
descriptions and the character by taxon matrix
used in this phylogenetic analysis have been
included as Appendices 1 and 2, respectively.
Added Taxa. The Begun et al. (1997a) matrix was first expanded to include five additional taxa (Turkanapithecus, Equatorius [sensu
Ward et al. 1999], Griphopithecus, Morotopithecus [sensu Gebo et al. 1997], and Ankarapithecus). Morphological data for these taxa were
obtained from the literature. Character coding
for the added taxa preferentially followed cladistic treatments of these taxa in other studies
by the authors of the Begun et al. (1997a) analysis to ensure consistency in character coding.
Additional character state information was
also incorporated from descriptions of holotypes and additional fossil material.
Morotopithecus was coded following Begun
and Güleç 1998 and Ward 1997a, and with descriptions of new postcranial fossils from the
Moroto II locality by Gebo et al. (1997) and
MacLatchy et al. (2000). Turkanapithecus was
coded following Rae 1997, Rose 1997, and
Ward 1997a. Additional character information
was obtained from Leakey et al. 1988. Follow-
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JOHN A. FINARELLI AND WILLIAM C. CLYDE
ing the separation of Equatorius from Kenyapithecus by Ward et al. (1999), coding of Equatorius followed the character coding for Kenyapithecus africanus by Rose (1997) and Ward
(1997a). Additional character information was
obtained from descriptions of fossil material
by Ward et al. (1999), Kelley et al. (2002), and
Sherwood et al. (2002). Begun (2000) has argued that Equatorius should be assigned to the
genus Griphopithecus on the basis of dental
similarity, although Kelley et al. (2002) argued
that differences between Equatorius and Turkish specimens attributed to Griphopithecus
support separation at the generic level. Recoding of the OTU Kenyapithecus after the removal
of Equatorius followed Rose 1997 and Begun et
al. 1997a for those characters attributable to K.
wickeri. Griphopithecus was coded following
Begun and Kordos 1997 and Begun and Güleç
1998, with additional character information
from Andrews et al. 1996, Alpagut et al. 1990,
and Begun 1992c. Ankarapithecus was coded
following Begun and Güleç 1998 and Alpagut
et al. 1996. In addition to the taxa added to the
analysis, newly described fossil material (Madar et al. 2002) provided character state information for the taxon Sivapithecus.
Added Characters. Five characters, which
have been proposed as phylogenetically informative but were not included in the Begun et
al. (1997a) analysis, were added to the data
matrix for this study. These were morphology
of the olecranon process and orientation of the
radial notch (Rose 1997), morphology of the
entocunieform facet on the navicular (Ward
1997a), torso shape (Schultz 1961; Ward
1997a), and a modification of the character
coding nasal aperture shape (Begun and Güleç 1998). All five characters were coded in
studies using the same coding scheme as Begun et al. (1997a), thus ensuring consistency.
Deleted Characters. The data matrix was
scanned for characters that were uninformative for the ingroup, and these were deleted
from the matrix. Following the numbering
scheme of Begun et al. (1997a) these were
characters 1, 7, 8, 9, 10, 11, 12, 13, 14, 15, 35,
36, 41, 46, 49, 50, 51, 64, 69, 70, 78, 97, 100, 161,
181, 225 and 240.
In several instances where multiple characters appeared to code for the same morpho-
logical trait, characters were consolidated to
avoid unduly weighting the impact of these
characters on the cladistic analysis. Combined
pairs (again following the numbering scheme
of Begun et al. 1997) include: characters 5 and
6 describing the morphology of the scapula,
18 and 19 describing the trochlea of the humerus, 39 and 40 describing the trapezoid facets, 47 and 48 describing ulnar articulation
with carpal elements, characters 94 and 95 describing the shape of the trochlea of the talus,
characters 143 and 188 describing the shape of
the nasal aperture (see above, ‘‘Added Characters’’), characters 213 and 215 describing the
morphology of the nasal clivus, and characters 232 and 233 describing the morphology of
the upper molars.
Character 88, describing the depth of the
femoral condyle, character 96, describing the
angle of the talar neck, and characters 112 and
113, describing the morphology of the first
metatarsal, were discarded from the matrix.
The femoral, tarsal, and metatarsal morphologies coded in these characters are believed to
represent derived hominin (5 human and all
direct ancestors [Tattersall et al. 1988]) characters associated with bipedalism and therefore are not homologous to the primitive propliopithecid condition (Jungers 1988; Ward
1997a). Additionally, characters 141, 142, and
176, describing the morphology of the maxillary sinuses, were excluded from the data matrix, as variation in maxillary sinus morphology in the hominoids has been demonstrated
to be a function of isometric scaling (Rae and
Kopp 2000).
Temporal Data
There is generally good stratigraphic control for hominoid fossil localities suggesting
that this information could be of potential value in resolving phylogenetic relationships
within this group. For this study, stratigraphic
data in the form of first and last appearance
events (FAEs and LAEs) for each hominoid
taxon were compiled from the literature.
Wherever available, radiometric ages or paleomagnetic data linking fossil localities to the
Geomagnetic Polarity Timescale (GPTS) of
Cande and Kent (1995) were used. However,
the temporal distributions of several taxa are
DATA CONGRUENCE IN HOMINOID PHYLOGENY
known only in relation to biostratigraphy
(e.g., Ouranopithecus), and for these cases appropriate regional faunal calibrations (e.g.,
European MN [Neogene Mammal] Zones)
were used to correlate these taxa into a universal temporal succession of first and last appearance events (FAEs and LAEs) for the
Hominoidea.
The coding of the stratigraphic character for
this study follows the methodology outlined
in previous stratocladistic analyses by Clyde
and Fisher (1997) and Bloch et al. (2001).
Range-through assumptions were made for
each taxon between its FAE and LAE, and
those taxa observed (or inferred by the rangethrough assumption) to exist at multiple levels
were assigned multiple character states (Clyde
and Fisher 1997). Although it is possible to
code sampling gaps in the stratigraphic character by skipping a letter or number in the
coding scheme (see Fisher 1992), a conservative coding scheme leaves such gaps uncoded
and treats intervals where no ingroup taxa are
observed as ‘‘no data’’ (Clyde and Fisher 1997;
Finarelli and Clyde 2002). Such intervals are
therefore not factored into the calculation of
stratigraphic parsimony debt.
Hominoids are observed to range from the
first appearance of Proconsul at Meswa Bridge
(biostratigraphically constrained to ca. 23.5
Ma [Pickford and Andrews 1981; Tassy and
Pickford 1983]) through the Recent. The stratigraphic data divide the range of the hominoids into 11 distinct stratigraphic intervals
(Fig. 1). The detailed stratigraphic information
that was used to order the FAEs and LAEs is
reported in Appendix 3. All of the boundaries
for stratigraphic character states are defined
by FAEs and/or LAEs, but not all appearance
events were used in defining character state
boundaries (e.g., Turkanapithecus; Fig. 1). That
is to say, in some cases the coding scheme was
coarsened such that the appearance events of
a particular taxon were subsumed within another stratigraphic character state, reflecting
areas where there is a lack of precision in the
stratigraphic data. Individual cases where this
occurred are discussed below.
Although the taxonomic assignment of the
Kisingiri and Tinderet material to the genus
Proconsul is rather secure, the potential FAE
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and LAE for this taxon is somewhat problematic. This is due to some uncertainty in attributing the Meswa Bridge (FAE) and Ngorora
Formation (LAE) material to Proconsul (see
Appendix 3). In the case of Meswa Bridge, if
the material were removed from the operational taxonomic unit of this study, the coding
of the stratigraphic character would not be altered as the point occurrence of Morotopithecus
would be coded with the Kisingiri and Tinderet levels, as the earliest stratigraphic horizon in the hominoid succession (‘‘a’’ in Fig. 1).
The LAE poses a more complicated problem.
With the assignment of the Ngorora Formation material to Proconsul, the young age of
these localities (approximately 12.5 Ma [Deino
et al. 1990; Hill et al. 1985, 2002]) creates a
large extension for Proconsul with the rangethrough assumption. However, if the Ngorora
material were not included in the OTU, then
the coding of the stratigraphic character for
Proconsul would comprise only a single character state (‘‘a’’ in Fig. 1). This could potentially have a considerable effect on the result
of the stratocladistic analysis. As such, we performed the stratocladistic analysis twice, using both coding schemes for Proconsul—first,
using a single stratigraphic character state (assuming that the Ngorora material is not Proconsul), and second, spanning multiple stratigraphic character states (assuming that it belongs to Proconsul—as depicted in Fig. 1). The
results of both analyses were identical. Therefore, the coding of the stratigraphic character
here reflects the most inclusive set of fossils attributed to Proconsul; however, it should be
noted that additional material (especially
from the Ngorora Formation) may significantly alter the appearance events for this taxon.
A recurring difficulty in coding the stratigraphic information for the hominoids is
‘‘point occurrences’’ in the fossil record. Turkanapithecus provides a good example. At Kalodirr in Kenya, Turkanapithecus is demonstrably younger than some of the Afropithecus material (Leakey and Leakey 1986b; Brochetto et
al. 1992), yet it is also found at the same level
as other Afropithecus fossils. It is easy to demonstrate contemporaneity of the two genera;
however, demonstrating that Afropithecus existed prior to, and exclusive of, Turkanapithecus
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JOHN A. FINARELLI AND WILLIAM C. CLYDE
FIGURE 1. Observed stratigraphic ranges for the hominoid taxa in this study. Stratigraphic ranges are based on
information presented in Appendix 3. Coding of the stratigraphic character for the Hominoidea is shown at the
right. The stratigraphic data for the ingroup produce 11 distinct stratigraphic intervals (coded ‘‘a’’ through ‘‘k’’).
First and last appearance events (FAEs and LAEs) define all stratigraphic character states. In several cases where
appearance events are poorly resolved, they are not used for defining character state boundaries. The stippled time
interval between the LAE of Oreopithecus and the FAE of Australopithecus is not coded in the stratigraphic character.
See text for discussion.
is difficult. Instead of coding two separate intervals for Afropithecus before the appearance
of Turkanapithecus and a geologically instantaneous interval comprising the overlap of Afropithecus and Turkanapithecus, they are conservatively coded as occupying a single stratigraphic level.
Similar arguments are made for Morotopithecus, Kenyapithecus, and Ankarapithecus. Morotopithecus is known from the Moroto fossil
sites in Uganda, which are radiometrically
dated to be older than 20.6 Ma (Gebo et al.
1997), and is known only from specimens at
this locality. If the coding of the stratigraphic
character were to coincide with all FAE and
LAE data, then one would have to code an interval before the occurrence of Morotopithecus,
an interval for the point occurrence of this taxon, and an interval following it. Instead, Morotopithecus is considered here to occupy the
stratigraphic interval defined by the FAE of
Proconsul and the FAE of Afropithecus. Kenyapithecus is known from Fort Ternan, and its
occurrence is entirely subsumed within the
upper range of Griphopithecus (Ward et al.
1999). Ankarapithecus is known from two localities in the Sinap Formation at Yassıören,
Turkey (Ozansoy 1965; Andrews and Tekkaya
1980; Alpagut et al. 1996) and correlates to the
interval defined by Sivapithecus and Dryopithecus. See Appendix 3 for details of the stratigraphic data.
The taxonomic assignment of the fossil material at Engelswies is uncertain. It is usually
DATA CONGRUENCE IN HOMINOID PHYLOGENY
assigned to ‘‘cf. Griphopithecus,’’ although
Heizmann and Begun (2001) urge caution and
suggest that the Engelswies hominoid may
represent a distinct genus. The FAE of Griphopithecus is therefore documented either at
Engelswies or at Paşalar (Heizmann and Begun 2001). Using either fossil locality for the
FAE does not alter the coding of the stratigraphic character (see Appendix 3). The lower
MN 5 FAE of Griphopithecus in Eurasia demonstrates that all of the western Kenyan Equatorius localities are contained within the range
of Griphopithecus. However, the lower limit on
this appearance event is less well constrained.
If the earliest possible calibration for the FAE
of Griphopithecus is used (Base MN 5 [see Heizmann and Begun 2001]) and the latest possible
calibration for the LAE of Afropithecus is also
used (Ad Dabtiyah is biostratigraphically correlated to the base of Maboko [see Gentry
1987a,b]), then some degree of overlap would
be implied, which would also imply a separate
stratigraphic character state coded on the basis two indefinite biostratigraphic correlations. Until either or both of these appearance
events are better constrained, the data do not
warrant the creation of this distinct character
state. As such, Afropithecus and Griphopithecus
are coded here as non-overlapping (Fig. 1).
The FAE of Sivapithecus is documented
within the Chron C5Ar.1, making it stratigraphically lower than the latest possible estimate for the LAE of Griphopithecus. A rangethrough assumption to the end of the MN 6
would place the LAE of Griphopithecus locally
at the Chron C5Ar.1r (Steininger et al. 1996).
However, coding an overlap in this case would
create a character state based on the correlation of the well-documented FAE of Sivapithecus to the terminus of a faunal zone for which
a range-through assumption must be made.
Conservatively, such an assumption of contemporaneity is not warranted. Similarly, Oreopithecus overlaps with the terminal portion
of the Sivapithecus range. The V2 horizon at
Baccinello is correlated to earliest MN 13
(Rook et al. 2000). If a similar range-through
assumption to the end of MN 13 were made,
Oreopithecus’s LAE would extend beyond Sivapithecus. Again, this correlation compares a
well-documented stratigraphic datum to the
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terminus of a faunal zone for which a rangethrough assumption is made, and coding of a
distinct character state is unwarranted.
Cladistic Analysis of the Hominoidea
The Morphologically Most Parsimonious
Cladograms
A cladistic analysis using the branch and
bound algorithm in PAUP* (version 4.0b10
[Swofford 2002]) recovered four most parsimonious cladograms with respect to the morphological data. Each had a tree length of 447
steps and a Retention Index (RI) of 0.69 (Fig.
2A). This value for the RI is somewhat low
when compared to other published RIs for
studies that include fossil taxa. For example,
Clyde and Fisher (1997: Table 2) compared RI
values across 29 studies of varied taxonomic
resolution. These studies had a median RI of
0.80, indicating that the degree of homoplasy
observed in the Hominoidea is somewhat
higher than for other taxonomic groups.
The strict consensus cladogram for the morphologically most parsimonious cladograms
(hereafter referred to as the MMPC) highlights topological features common to all of
the cladograms as well as areas of ambiguity
(Fig. 2B). Proconsul is the sister taxon to all
other ingroup taxa. Unfortunately, this does
not resolve the debate over the phyletic position of Proconsul, because this position is consistent with its being either a basal hominoid
(Andrews and Martin 1987a; Andrews 1992)
or an undifferentiated Miocene catarrhine
(Harrison 1987; Harrison and Sanders 1999).
The most notable feature of the MMPC is the
distinction between a postcranially primitive
clade of early to middle Miocene hominoids
from East Africa and a postcranially derived
hominoid clade that includes both late Miocene Eurasian forms and all of the extant hominoid genera. Morotopithecus is joined to the
base of the derived clade, supporting MacLatchy et al. (2000), who hypothesized the existence of two evolutionarily distinct hominoid lineages in East Africa by 20 Ma on the
basis of several derived features of the femur
and axial skeleton. However, it is not the inclusion of Morotopithecus in this phylogenetic
analysis that is responsible for the division of
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JOHN A. FINARELLI AND WILLIAM C. CLYDE
FIGURE 2. A, The four morphologically most parsimonious cladograms (MMPC) recovered for the character by
taxon matrix compiled for the 18 ingroup taxa (Appendix 2). Each cladogram has a tree length of 447 steps and an
RI of 0.69. B, The strict consensus cladogram of the MMPC. The morphologically most parsimonious hypotheses
distinguish two distinct clades of hominoids, a clade of ‘‘archaic’’ hominoids from the early Miocene of East Africa,
and a clade of ‘‘modern’’ hominoids, including all extant hominoids and the early Miocene hominoid Morotopithecus.
Numbers above branches indicate Bremer Decay indices for corresponding internal nodes.
DATA CONGRUENCE IN HOMINOID PHYLOGENY
the hominoids into these two separate clades.
The phylogenetic analysis was repeated on the
data set while excluding Morotopithecus, and a
set of eight cladograms was recovered, each
distinguishing a monophyletic radiation of
early to middle Miocene hominoids seen in
the MMPC. The topologies of each of these
eight cladograms were identical to the
MMPC, except that the relative position of
Proconsul varied across the set creating a polytomy between Proconsul and the two hominoid clades. Thus, early Miocene hominoids
from East Africa form a distinct clade in the
MMPC, and the early appearing Morotopithecus is allied with the postcranially derived
clade of hominoids.
This hypothesis parallels, although is not
exactly identical to, Pilbeam’s (1997) concept
of distinct evolutionary histories for hominoids of ‘‘archaic’’ and ‘‘modern’’ aspects. For
ease of reference, Pilbeam’s terminology will
be adopted here, and the clade of postcranially primitive hominoids from the early and
middle Miocene of East Africa (Afropithecus,
Turkanapithecus, Equatorius, Kenyapithecus, and
Griphopithecus) will be informally referred to
as ‘‘archaic’’ hominoids, and the postcranially
derived clade (Morotopithecus, etc.) as ‘‘modern’’ hominoids (Fig. 2B). Analysis of unambiguous character state reconstructions (DELTRAN optimization) demonstrates that the
clade of ‘‘archaic’’ hominoids is supported by
synapomorphies primarily centered on the
morphology of the face (Table 1). Derived features of the vertebrae and hindlimb, which are
thought to be associated with positional behavior and locomotion (Ward 1997a), are reconstructed as synapomorphies for the clade
of ‘‘modern’’ hominoids (Table 1).
Within the clade of ‘‘modern’’ hominoids,
there is a monophyletic clade of great apes
(Hominidae), which includes distinct clades of
South Asian great apes (Ponginae: Ankarapithecus, Sivapithecus, Lufengpithecus, and Pongo)
and African great apes (Homininae: Pan, Gorilla, and Australopithecus) (Fig. 2B). The interrelationships between the pongines, the hominines, and the late Miocene European hominoid Ouranopithecus are ambiguous in the
strict consensus, as Ouranopithecus is alternately hypothesized as the sister taxon to the
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hominines (Andrews 1992; Andrews et al.
1996; Dean and Delson 1992), to the pongines
(Schwartz 1990; Moyà-Solà and Köhler 1993),
or to all extant great apes (Fig. 2A). However,
the Ouranopithecus/hominine/pongine clade
is consistently derived in the MMPC with respect to Dryopithecus (contra Begun 1992b,
1994; Begun et al. 1997a; Begun and Güleç
1998). Additionally, the interrelationships
among the chimpanzee, gorilla, and human
lineages are not resolved in the strict consensus. A large suite of synapomorphies including postcranial, cranial, and dental characters
unites the great ape clade (Table 1). Synapomorphies associated with their unique facial
morphology and extreme heteromorphy of
the upper incisors unite the pongine clade,
whereas morphology primarily associated
with the hands and feet (e.g., fusion of os centrale) and cranium (e.g., increased robusticity
of the supraorbital torus, broad supraorbital
sulcus) unite the hominine clade (Table 1).
The MMPC have tree lengths of 447 steps
with 201 homoplasies. However, 17 cladograms were recovered with tree lengths of 448
steps (202 homoplasies) and 67 cladograms
with 449 steps (203 homoplasies). By 455 steps
(209 homoplasies) there are more than 16,000
recovered cladograms. This large number of
cladograms with only slightly higher tree
lengths raises concern over the strength of
support for the topologies observed in the
strict consensus cladogram. Although this is
certainly not a new debate, there is still no
method to determine if the cladograms with
201 homoplasies are significantly better than
those with 202 homoplasies (Wagner 1995). In
the absence of a rigorous criterion for determining significance, we used additional analyses to evaluate the strength of support for the
phylogenetic statements in the most parsimonious cladograms.
Consensus Analyses
By evaluating strict consensus cladograms
for successively higher tree lengths, the number of additional steps needed to ‘‘break’’ a
node can be used as a measure of the relative
strength of support for the phylogenetic statements in a cladogram (Bremer 1988). Bremer
Decay indices were calculated for the strict
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JOHN A. FINARELLI AND WILLIAM C. CLYDE
TABLE 1. Unambiguous synapomorphies uniting major clades in the MMPC: character states reconstructed using
the DELTRAN optimization.
Clade
‘‘Hominoids of Archaic Aspect’’
‘‘Hominoids of Modern Aspect’’
Hominidae
Ponginae
Homininae
Description
Reduction of molar cusps and lower canine
Deepening of the zygomatic and more vertical orientation of zygomatic
Lowering of the nasal aperture relative to the orbits
Increased length of nasal bones
Deep canine fossa
Lengthening of nasoalveolar clivus
Increased vertebral body height
More dorsally positioned transverse process
Increased asymmetry of the femoral condyle
More medially asymmetric trochlear keel
Loss of articulation of ulna with pisiform/triquetral
Broadening of sternebrae
Increased size of plantar tubercle
Increased cuboid wedging
Loss of prehallux facet (MT 1)
Increased robusticity of MT 2 and MT 5
Shift in the foot axis through the second digit
Reduction of male canine size
Upper P3: reduced cusp heteromorphy, reduced paracone
Lower P3 metaconid present
Lengthening of lower P4
Loss of metacones, increase in upper molar crown size
Relative increase in lower M1 size, lengthening of upper M1 and M2
Increased heteromorphy of the lateral incisors
Inflation of glabella
Presence of supraorbital torus and superciliary ridges
Reduction of frontal sinus size
Increased length of neurocranium
Reduction of articular tubercle and articular/tympanic
fused to temporal
Raising of nasal aperture relative to alveolar plane
Presence of deep incisive fossa
Lengthening of nasoalveolar clivus
Large incisive canal present
Greatly increased lateral incisor heteromorphy
Greatly increased orbital breadth
Zygomatics positioned more anteriorly and vertically
oriented
Decreased distance from nasal aperture to orbits
(primitive reversal)
Deep canine fossa
Piriform nasal aperture
Fused os centrale
Concavoconvex centrale facet on capitate
Angled posterior talar facet
Decreased cuboid peg
Increase in size of phalangeal flexor ridges
Longer upper P4
Increased robusticity of supraorbital torus, broad supraorbital sulcus
Inferior orientation of the nuchal plane
More horizontal orientation of the zygomatics
Inflated maxillary alveolar process
Larger lesser palatine foramina
Pyramidal process inferiorly broad
Pterygoid process compressed
Characters
111, 117
161, 169
171
174
179
197
57
59
70
15
36
60
80
83
90
91, 95
94
100
103, 104
107
108
110, 131
119, 128
121
143
144, 145
148
155
157, 158
172
195
197
199
121
141
162, 169
164
179
200
28
42
78
82
97
124
144, 146
154
169
182
186
190
191
DATA CONGRUENCE IN HOMINOID PHYLOGENY
consensus of the MMPC using the program
AutoDecay (Version 5.0 [Eriksson 2001]) and
PAUP* (heuristic searches with 100 random
sequence additions) (Fig. 2B). The Bremer Decay indices calculated for the MMPC are very
low; no internal node in the strict consensus
persists past three additional steps. A distinct
clade of ‘‘modern’’ hominoids is recoverable
only through two additional steps, and the
‘‘archaic’’ clade is recoverable through a single
additional step. Several clades are not recoverable through even one additional step (Bremer Decay Index 5 0; Fig. 2B), such as the African great apes, as one of the cladograms
with a tree length of 448 allies Ouranopithecus
with Australopithecus. This low Bremer support is likely the result of the relatively high
proportion of homoplasy and highly incomplete preservation for several of the fossil taxa.
However, strict consensus cladograms are
highly insensitive to repeated clustering of
taxa when the phylogenetic positions of a few
taxa in the ingroup are highly mobile (Wilkinson 1999), and such mobility can be greatly
enhanced by incomplete representation in the
data matrix, and therefore highly incomplete
fossil taxa. Because of this insensitivity, relationships preserved in strict consensus cladograms can be inferred as strong statements,
but important phylogenetic statements can be
lost when relying solely upon strict consensus
methods (Wilkinson 1994, 1996). We therefore
used reduced consensus analysis (Wilkinson
1994, 1996, 1999) to uncover any such obscured statements, highlighting groupings
that are consistently implied by a set of cladograms if one or more taxa with variable positions are removed from consideration. It is
important to note that this method does not
ignore data. The entire data matrix is used to
recover the set of cladograms; taxa are only
subsequently pruned to uncover repeated topologies lost in the strict consensus (Wilkinson 1994, 1999).
For the reduced consensus analyses, we
used the program ‘‘strict.exe’’ in the REDCON
software package (Version 3.0 [Wilkinson
2001]). Reduced consensus analyses demonstrate that the morphological support for the
‘‘archaic’’ lineage is limited. These analyses
were only able to recover parts of the ‘‘archa-
623
ic’’ hominoid clade upon exclusion of mutually segregating units (i.e., if Turkanapithecus is
excluded, then a Kenyapithecus/Equatorius/
Griphopithecus clade is recovered; if Kenyapithecus/Equatorius/Griphopithecus are excluded,
then Turkanapithecus/Afropithecus is recovered
through three additional steps, and through
four steps if Kenyapithecus is removed.
From the reduced consensus analysis, it
was apparent that much of the poor nodal
support was due to Kenyapithecus, which was
repeatedly excluded from reduced consensus
partitions at and beyond three additional
steps. The great ape clade is supported in reduced consensus through four additional
steps upon removal of only Kenyapithecus, and
it can be recovered through six additional
steps if Lufengpithecus is also excluded. When
Kenyapithecus is excluded, a clade uniting Pongo and Sivapithecus is recovered through six
additional steps. Although these results do
imply a good deal of ambiguity in the relationships of late Miocene European apes and
the hominines, they argue for distinct hominine and pongine lineages. In addition, the repeated exclusion of Kenyapithecus from reduced consensus cladograms demonstrates
that the phylogenetic position of this taxon is
highly unstable at slightly longer tree lengths.
It is likely that this instability is due to incomplete preservation. Kenyapithecus is coded for
only 25% of the characters in the data matrix
(Appendix 2).
Bootstrap Analysis
Bootstrapping was also used to evaluate
support for MMPC topologies (Felsenstein
1985). Two hundred characters were randomly resampled with replacement for 1000 heuristic search replicates (ten random sequence
additions per replicate) and a majority-rule
consensus cladogram was produced. Most of
the nodes observed in the MMPC are left ambiguous in the 50% majority-rule cladogram
for the bootstrap (Fig. 3A). Additionally, those
nodes that were resolved have low bootstrap
proportions, indicating only marginal support. It is likely that this, too, is a function of
the incompleteness of certain taxa in the data
matrix, which increases ambiguity under resampling techniques such as the bootstrap. We
624
JOHN A. FINARELLI AND WILLIAM C. CLYDE
FIGURE 3. A, The majority-rule bootstrap consensus of the full character by taxon matrix (heuristic search ten random sequence additions; 1000 replicates). Many of the internal nodes in the bootstrap topology are ambiguous,
and bootstrap proportions are generally low. These ambiguities are introduced by the highly incomplete taxon
Kenyapithecus. B, The majority-rule bootstrap consensus for the same character data, excluding Kenyapithecus. Numbers above branches indicate bootstrap proportions for corresponding internal nodes. Bootstrap proportions for
the internal nodes resolved in A are either maintained or increased in B.
DATA CONGRUENCE IN HOMINOID PHYLOGENY
noted above that Kenyapithecus played a large
role in the low Bremer support for many of the
nodes in the MMPC. Therefore, a second bootstrap was performed on the data set excluding
Kenyapithecus (Fig. 3B). The topology of this
second bootstrap is fully consistent with that
of the first, only more resolved, and the bootstrap proportions indicate stronger support
for several of the internal nodes. Upon removal of Kenyapithecus, an Equatorius/Griphopithecus clade is also recovered. Removal of Kenyapithecus also serves to separate the clade of
‘‘modern’’ hominoids (albeit without Morotopithecus) relative to early Miocene hominoids
from East Africa with strong support, as well
as increasing the support for a distinct clade
of great apes, including Oreopithecus.
The bootstrap topologies do differ significantly from the strict consensus cladogram in
several respects. In the strict consensus of the
MMPC, both the pongine clade and Ouranopithecus are hypothesized as derived relative
to Dryopithecus and Oreopithecus. In both bootstraps (with and without Kenyapithecus), all
European hominoids are joined in an unresolved polytomy with the hominines (Fig.
3A,B). The support for this group as a clade
that is both distinct from and derived in relation to the South Asian hominoids is extremely weak (52% in both bootstraps), indicating that the relative positions of the Homininae and Ponginae with respect to late Miocene European hominoids is rather weakly
supported by the morphological data, despite
its apparently clear resolution in the MMPC.
The polytomy in the bootstrap topologies
formed by the South Asian hominoids does
distinguish a Pongo/Sivapithecus clade with
reasonable support. The lack of resolution of
a monophyletic Ponginae here is likely also a
function of incompleteness of Ankarapithecus
and Lufengpithecus. These taxa are 46% and
38% coded, respectively, whereas Sivapithecus
and Pongo are 72% and 100% coded, respectively. As the incompleteness of a taxon increases, it becomes more likely that resampling techniques, such as the bootstrap, will
sample heavily from missing character data
for that taxon. This leads to an increase in the
ambiguity of its placement within and among
bootstrap replicates, and serves to lower boot-
625
strap proportions. It is likely that much of the
uncertainty in the position of Lufengpithecus
and Ankarapithecus is caused by the relatively
poor preservation of a few key taxa.
Two additional bootstraps were performed
removing either Ankarapithecus or Lufengpithecus in addition to Kenyapithecus. The bootstrap
topologies produced for the remaining taxa
for these additional analyses were identical to
the bootstrap excluding Kenyapithecus (Fig. 3B)
with several notable exceptions. The removal
of the more incomplete Lufengpithecus causes
the bootstrap proportion for Sivapithecus/Pongo to rise to 79%, and a monophyletic Ponginae is recovered with 73% support. However,
if Ankarapithecus is excluded and Lufengpithecus retained, then the bootstrap proportion for
Sivapithecus/Pongo rises to 88%, and a monophyletic Ponginae is recovered 52% of the
time. The removal of additional incompletely
coded taxa decreases some of the ambiguity in
the bootstrap results in other regions of the
cladogram. The bootstrap proportions for the
Hominidae increase in both of these analyses
to about 90%, and there is an increase support
for the Equatorius/Griphopithecus clade (approximately 70%), and a weakly supported
(about 55%) clade of ‘‘modern’’ hominoids is
recovered. None of the bootstrap analyses recovered a distinct clade of ‘‘archaic’’ hominoids.
Judging from these various analyses, it is
clear that poorly preserved fossil taxa and relatively high levels of homoplasy infuse considerable uncertainty into the cladistic analysis of hominoids. In fact only a few partitions
of the MMPC (e.g., Homininae, Sivapithecus/
Pongo clade) are consistently supported and
the results are highly sensitive to the choice of
ingroup taxa.
Homoplasy Partitioning
To understand better how homoplasy is distributed across the hominoid clade, we performed additional character analyses. The ingroup was divided into two subsets of taxa
based upon the split between the ‘‘archaic’’
and ‘‘modern’’ hominoid clades observed in
the MMPC and reevaluated to examine the
partitioning of homoplasy across the ingroup.
The ‘‘archaic’’ group here included Proconsul
626
JOHN A. FINARELLI AND WILLIAM C. CLYDE
with Afropithecus, Turkanapithecus, Equatorius,
Kenyapithecus, and Griphopithecus. The ‘‘modern’’ group was composed of the remainder of
the ingroup taxa. The ‘‘modern’’ hominoid
subset produced eight most parsimonious
cladograms. The topology of their strict consensus is identical to the analogous partition
of the strict consensus of the MMPC, except
Morotopithecus and Hylobates form a polytomy
at the base of the consensus cladogram. A single most parsimonious cladogram was recovered for the ‘‘archaic’’ subset with a topology
identical to that seen in all of the MMPC.
Interestingly, the degree of observed homoplasy is markedly different between these
taxonomic partitions. The RI for the ‘‘modern’’
hominoid clade is 0.54, whereas the RI for the
‘‘archaic’’ hominoid clade is 0.90. The morphological data for the clade of ‘‘archaic’’
hominoids is therefore internally consistent,
indicating that the resolution problem noted
in the consensus and bootstrap analyses for
these taxa is not the result of excessive homoplasy, but rather of highly incomplete preservation (with the exception of Proconsul). The
elevated rate of homoplasy for the Hominoidea in general is apparently confined almost
entirely to the clade of ‘‘modern’’ hominoids.
We further examined homoplasy across
dental and postcranial character partitions to
determine the relative proportion of homoplasy in these anatomical regions. Average character RI values obtained for the set of postcranial skeletal characters (1–97) and dental characters (characters 98–134) were calculated
across the strict consensus of the MMPC. Results show that postcranial characters (mean
RI 5 0.75) exhibit lower levels of homoplasy
than dental characters (mean RI 5 0.59). However, several aspects of the dental character
subset raise concern about simply accepting
this pattern as a reflection of the degree of homoplasy in the hominoid dentition and postcrania. First, the dental character subset is
characterized by more complete preservation.
The dental subset is 90% coded in the data
matrix, as compared with only 49% for postcranial characters. Second, the dental character data are represented by a smaller number
of characters in the data matrix. Both of these
factors may influence the amount of homopla-
sy observed when comparing the fit of these
character partitions to the MMPC. For instance, convergence may be masked by the relatively poorer preservation of postcranial
characters, or the larger number of postcranial
characters may implicitly weight the outcome
of a morphology-based analysis, generating a
cladogram that is fundamentally biased in favor of the postcrania.
To eliminate these potential sources of bias,
a weighted rarefaction analysis was performed. Dental character data were randomly
deleted from the fossil taxa in the original data
matrix until the completeness (proportion of
cells coded in the data matrix) equaled that of
the postcranial character data (29% across fossil taxa). This procedure was repeated 100
times creating a set of replicate data sets that
exhibit equal preservation in both the dental
and postcranial character partitions for the
fossil taxa in this analysis. Weighting the dental characters by 2.62 equalized the relative input of the postcranial and dental character
partitions (97 vs. 37 characters, respectively)
in determining the resulting most parsimonious cladogram for each replicate. The set of
optimal cladograms was then recovered for
each rarefaction with the branch-and-bound
algorithm, allowing a null hypothesis (H0: no
significant difference in level of homoplasy
between dental and postcranial characters exists) to be tested.
Mean character RI scores for the dental and
postcranial character blocks were calculated
separately over the returned cladograms for
each rarefaction of the dental data. The distribution of rarefaction mean character RIs for
postcranial characters is centered on an average of 0.78, compared with a distribution
mean of 0.73 for average dental character RI
scores (Fig. 4 top). A paired t-test on the rarefaction data demonstrates that postcranial RI
scores are significantly higher than the corresponding RI scores for dental characters (t 5
5.965, p K 0.01; one-tailed). Examination of
each rarefaction on a case-by-case basis reveals that 72% of the cases are characterized
by higher mean postcranial RI values (Fig. 4
bottom).
These results support earlier studies on
hominoid data sets, which concluded that
DATA CONGRUENCE IN HOMINOID PHYLOGENY
627
FIGURE 4. Top, Histogram of RI values for postcranial and dental characters after rarefaction. The rarefaction analysis controlled for differential preservation and implicit weighting. Arrows plot mean values for each distribution.
The mean for the postcranial character distribution is significantly higher than the mean for the dental characters
(Paired t-test: t 5 5.965; p K 0.01). Bottom, Case-by-case comparison of each rarefaction showing higher RI values
for the dental characters than the postcranial characters for the majority of cases (72/100).
dental character data may not reliably recover
phylogenetic relationships (Hartman 1988;
Collard and Wood 2000). However, these earlier approaches compared the performance of
anatomical subsets in reproducing the cladistic topologies recovered in molecular studies,
and therefore assumed the correctness of the
molecular hypothesis. Regardless of any argument for preferring a cladogram derived
from molecular evidence (Collard and Wood
2000: p. 5003), it must remain an assumption
that the molecular data recovered ‘‘true’’ hom-
628
JOHN A. FINARELLI AND WILLIAM C. CLYDE
TABLE 2.
Total parsimony scores for the phylogenetic hypotheses proposed in this study.
Phylogenetic hypothesis
MMPC 1
MMPC 2
MMPC 3
MMPC 4
Stratocladistic hypothesis
Morphologic
tree length
Stratigraphic
parsimony debt
Stratigraphically
augmented tree length
RImorph
RIstrat
447
447
447
447
455
37
35
37
38
25
484
482
484
485
480
0.69
0.69
0.69
0.69
0.68
0.61
0.63
0.61
0.60
0.73
inoid phylogenetic relationships. Thus, these
studies are actually comparing the goodnessof-fit of various anatomical character partitions with a molecular topology (or the congruence between the molecular and various
morphological data partitions). This approach
is different in that the two anatomical subsets
are equalized with rarefaction and weighting
and then evaluated relative to one another using a phylogenetic hypothesis that is derived
from their combined input. This analysis provides a quantitative measure of relative performance of postcranial and dental character
data subsets under the parsimony criterion
and suggests that, when using this optimization criterion, postcranial characters are more
reliable phylogenetic indicators in the Hominoidea.
Incorporation of Stratigraphic Data
Stratigraphic Debt in the Morphologically
Most Parsimonious Cladograms
The coded stratigraphic character was appended to the morphological data matrix and
stratigraphically augmented tree lengths (i.e.,
morphological tree length 1 stratigraphic
parsimony debt) were calculated for the
MMPC. The stratigraphically augmented tree
lengths, stratigraphic debt values, and the RI
scores relative to the stratigraphic character
(RIstrat [Clyde and Fisher 1997]) for each of the
MMPCs have been compiled in Table 2.
The maximum possible stratigraphic debt
value for the observed stratigraphic ranges is
94 steps and the minimum possible value is 0,
corresponding to an anagenetic lineage in
which the ingroup taxa evolve in the order of
their first appearances. It should be noted that
because the stratigraphic character evaluates
all hypotheses as phylogenetic trees rather
than as cladograms, it is always possible (given the coding scheme for the stratigraphic
character used here) to hypothesize an anagenetic lineage without ghost ranges (Clyde
and Fisher 1997). Under stratocladistics, when
cladograms (such as the MMPC) are evaluated
with respect to the stratigraphic character,
they are actually treated as their isomorphic
phylogenetic trees (identical branching pattern and no observed ancestors). Therefore,
when evaluating stratigraphically augmented
tree lengths, a cladogram must have some
non-zero stratigraphic debt value associated
with the ranges connecting taxa at terminal
nodes to hypothetical ancestors at internal
nodes (Finarelli and Clyde 2002). The RIstrat
values for the MMPC (0.60 to 0.63; Table 2) are
higher than the median stratigraphic debt value (0.53) reported in Clyde and Fisher (1997:
Table 2) across 29 cladistic analyses in the literature. Thus, the hominoids show a better
than average congruence between the stratigraphic order of observed appearance events
and cladistic branching order.
Partitioning of Stratigraphic Parsimony Debt
In order to evaluate the partitioning of
stratigraphic parsimony debt, the ingroup
was again divided into the ‘‘archaic’’ and
‘‘modern’’ hominoid subsets and the stratigraphic character was recoded for each subset
independently. The RIstrat of the most parsimonious cladogram for the ‘‘archaic’’ subset is
0.67, indicating a much better fit of the stratigraphic data to this part of the phylogeny than
to the strict consensus for the cladograms recovered for the ‘‘modern’’ subset (RIstrat 5
0.38). This difference in RIstrat values mirrors
the pattern observed for morphology, indicating that both the morphological and strati-
DATA CONGRUENCE IN HOMINOID PHYLOGENY
graphic data for the ‘‘modern’’ hominoids display a poorer fit to the proposed cladistic relationships.
This poorer fit of stratigraphic data to the
cladistic ordering of the ‘‘modern’’ hominoids
is largely due to long ghost lineages for Hylobates and Pongo caused by their essentially
nonexistent fossil records. Another problem
lies in the early appearance of Sivapithecus.
The relatively primitive morphology of Hylobates requires a ghost lineage extending back
to the FAE of Sivapithecus. In addition, significant accumulation of stratigraphic debt is incurred by the implied close relationships of
Ankarapithecus, Lufengpithecus, and especially
Pongo. Because of the active geological setting
of the Siwaliks, it is possible that the observed
FAE of Sivapithecus more closely approximates
its true FAE than is the case for coeval hominoid taxa. Counterintuitively then, a higher
preservation probability for Sivapithecus, as
compared with its hypothesized close relatives in the MMPC, may contribute to the low
RIstrat observed in the ‘‘modern’’ hominoids. If
this is true then the assumption of uniform
preservation probabilities (Fisher 1992; Rieppel and Grande 1994) may be violated for
some of the stratigraphic levels defined above.
However, to test this rigorously would require
incorporating quantitative range extensions
that require significantly more stratigraphic
information than is presently available.
Stratocladistic Analysis of the Hominoidea
For the stratocladistic analysis, the set of all
cladograms with tree lengths below a threshold value was imported into MacClade (version 3.08a [Maddison and Maddison 1999]).
As there is currently no automated search algorithm for performing a stratocladistic analysis, this threshold value was chosen on the
basis of the number of returned cladograms,
seeking to maintain manageability of the returned set. The imported trees were evaluated
relative to total parsimony debt (morphological 1 stratigraphic parsimony debt). Manual
branch swapping was performed to uncover
additional topologies with lower total debt,
and hypotheses of explicit ancestry were tested (Fisher 1992; Clyde and Fisher 1997; Bloch
et al. 2001). Although it is theoretically pos-
629
sible to guarantee the return of the most parsimonious phylogenetic tree in a stratocladistic analysis using the ‘‘debt ceiling’’ approach
of Fisher (1992), in practice this is usually not
feasible because of the large number of potential hypotheses and the lack of an automated
algorithm. The manual heuristic search of topologies and ancestor configurations is not
guaranteed to uncover the most parsimonious
tree(s). However, for a relatively small number
of taxa such as in this analysis, a researcher
can reliably minimize total parsimony debt
with the manual search in MacClade (Fisher
1992).
Following the methodology detailed above,
an arbitrary cutoff of 455 steps was used in a
branch-and-bound search, recovering 28,093
cladograms. Because the stratigraphic data do
fit the MMPC relatively well and the number
of stratigraphic character states is small relative to the total number of morphological
character states, reductions of stratigraphic
debt are minor compared with the corresponding morphological debt that is incurred.
Evaluation by manual branch swapping and
incorporation of explicit statements of ancestry recovered a single most parsimonious
phylogenetic tree (Fig. 5). This tree has a stratigraphically augmented tree length of 480
steps, a morphological tree length of 455 steps
(RImorph 5 0.68), and 25 units of stratigraphic
parsimony debt (RIstrat 5 0.73; see Table 2). As
in Clyde and Fisher (1997), a substantial increase in the value of RIstrat was observed,
without a corresponding decrease in RImorph.
This stratocladistic hypothesis shows a primarily pectinate succession of hominoids
through the early Miocene (Fig. 5). Kenyapithecus is hypothesized as the ancestor to all extant hominoids and their fossil relatives (Andrews and Martin 1987a). This statement of
ancestry involves no increase in the morphological debt for the hypothesis. Rather it simply reduces the stratigraphic debt relative to
the corresponding hypothesis without an explicitly hypothesized ancestor. This implies
that no autapomorphic character-state transitions occur in Kenyapithecus, which is likely a
function of the original character coding (see
Bloch et al. 2001). In addition, this proposed
ancestral position of Kenyapithecus is at odds
630
JOHN A. FINARELLI AND WILLIAM C. CLYDE
FIGURE 5. The most parsimonious phylogenetic tree recovered in the stratocladistic analysis. The stratigraphically
augmented tree length of this stratocladistic hypothesis is 480 steps, with 25 steps of stratigraphic parsimony debt
(RImorph 5 0.68, RIstrat 5 0.73). Kenyapithecus is recognized as an ancestral taxon in this hypothesis.
with recent phylogenetic analyses, which generally consider Kenyapithecus as too primitive
to be a close relative of the extant hominoids
(Begun et al. 1997a; McCrossin and Benefit
1997), although these studies did incorporate
‘‘Kenyapithecus’’ prior to its partition into the
two OTUs, Equatorius and Kenyapithecus, used
in this study (Ward et al. 1999). Given the incompleteness of coding for Kenyapithecus, support for this explicit statement of ancestry is
rather weak. However, it is important to note
that the array of ‘‘archaic’’ taxa in a pectinate
manner is strongly supported by the stratigraphic data, as it minimizes the amount of
stratigraphic debt incurred by the hypothesis
by eliminating the long ghost lineage implied
by Morotopithecus in the MMPC.
Comparison of Morphological and
Stratocladistic Hypotheses
As stratocladistic hypotheses are phylogenetic trees, it is desirable to compare the topologies of the MMPC with the cladistic topology that is consistent with the phylogenetic tree recovered by the stratocladistic analysis. Such a topological comparison between
the cladistic and stratocladistic hypotheses reveals several important differences (Fig. 6).
Most significantly, the division of the Hominoidea into ‘‘archaic’’ and ‘‘modern’’ clades is
not recovered in the stratocladistic analysis.
Although Afropithecus and Turkanapithecus remain sister taxa, the remainder of the ‘‘archaic’’ taxa are arranged in pectinate succession
DATA CONGRUENCE IN HOMINOID PHYLOGENY
631
FIGURE 6. Comparison of the MMPC (left) and the cladogram associated with the stratocladistic hypothesis. Although Afropithecus and Turkanapithecus remain grouped as sister taxa in the stratocladistic hypothesis, the remainder of the ‘‘archaic’’ clade is not recognized. These taxa are arranged in a pectinate manner, eliminating the
long ghost lineage implied by the position of Morotopithecus in the MMPC. Two arrows point to taxa whose relative
positions in the cladogram are significantly changed. Morotopithecus (an early-appearing hominoid with derived
postcranial morphology) is displaced baseward in the stratocladistic hypothesis, whereas the late-appearing taxon
(Oreopithecus) is displaced crownward. See text for discussion.
following their order of appearance, essentially grafting the ‘‘archaic’’ hominoids into
the implied gap between the appearances of
Morotopithecus and Sivapithecus.
The displacement of the early-appearing
Morotopithecus baseward in the stratocladistic
hypothesis is accompanied by the crownward
movement of the late-appearing Oreopithecus.
The savings in stratigraphic parsimony debt
associated with these movements are obvious.
In the case of Morotopithecus (27.5% coded in
the data matrix), incomplete coding of morphology makes its position relatively poorly
constrained. Although incomplete fossil preservation facilitates uncertainty in the phylogenetic position of some taxa it is not a necessary condition for it. For instance, the displacement of Oreopithecus (in this case crownward) eliminates the stratigraphic debt
associated with its long ghost lineage in the
MMPC (Fig. 7D). Unlike Morotopithecus however, Oreopithecus is coded for 70% of the char-
acters in the data matrix. Closer examination
revealed that the repositioning of Morotopithecus baseward and the inverted relative positions of Dryopithecus and Ouranopithecus facilitated the repositioning of Oreopithecus in the
stratocladistic hypothesis. Without both of
these movements, displacing Oreopithecus
crownward is associated with large increases
in morphological debt that are not compensated for by the decreases in the stratigraphic
debt. Therefore, the position of other taxa including Morotopithecus has a large impact on
the pattern of character polarity, making the
switch in the relative position of Ouranopithecus and Dryopithecus possible, which in turn
allows the crownward movement of Oreopithecus to dramatically lower stratigraphic
debt.
The change in the relative position of Dryopithecus and Ouranopithecus highlights another significant difference between the MMPC
and the stratocladistic hypothesis—the pon-
632
JOHN A. FINARELLI AND WILLIAM C. CLYDE
FIGURE 7. A, Graphic representation of the stratigraphic debt incurred upon the strict consensus of the MMPC. B,
Graphic representation of the stratigraphic debt incurred upon the stratocladistic hypothesis. Note that Kenyapithecus is recognized as an ancestral taxon in this hypothesis. As in Figure 1, the stippled region represents a sampling gap that is not coded in the stratigraphic character. C, Arrow points to the removal of large amounts of stratigraphic debt by displacing Morotopithecus baseward in the stratocladistic hypothesis and thus eliminating distinct
DATA CONGRUENCE IN HOMINOID PHYLOGENY
gines. Associated with the repositioning of
these two European hominoids is the displacement of the pongine clade, such that Dryopithecus is consistently derived relative to the
South Asian hominoids (Fig. 6). The stratocladistic hypothesis produced a topology similar to the results of a cladistic analysis performed by Begun (2002) in this respect. However, in that analysis Lufengpithecus and Ankarapithecus remained in a monophyletic
pongine clade with Pongo and Sivapithecus (as
in the MMPC of this study). In contrast, although Sivapithecus and Pongo remain united
as sister taxa, the positions of both Lufengpithecus and Ankarapithecus change, placing
them as sister taxa to European and African
hominoids in the stratocladistic hypothesis
(Fig. 6). Unfortunately, Begun (2002) only included Griphopithecus and Afropithecus from
the ‘‘archaic’’ clade, and therefore that analysis does not address the question of support
for distinct early Miocene hominoid lineages.
It is important to note here that the stratocladistic hypothesis is not simply arranging
the ingroup taxa by their order of appearance.
For instance, the relative positions of Pongo
and Hylobates are not altered between the two
hypotheses (Figs. 6, 7) even though these hypothesized positions imply long ghost lineages. Repositioning either taxon crownward
significantly lowers stratigraphic debt, but
any such movement simultaneously creates
large amounts of morphological debt, and the
total parsimony criterion rejects these hypotheses.
The topological differences between the cladistic and stratocladistic hypotheses highlight
two intervals in the fossil record of the Hominoidea where significant incongruence exists
between the morphological and temporal
data. The first interval occurs in the early Mio-
633
cene of East Africa. The position of Morotopithecus as the sister taxon to all extant and late
middle to late Miocene Eurasian hominoids in
the MMPC implies a large gap in this group’s
fossil record spanning the early to middle
Miocene of East Africa. Numerous hominoid
localities exist in this region that span this period of time (approximately 21–14 Ma). The
long ghost lineage in the morphological cladogram is eliminated in the stratocladistic hypothesis by baseward displacement of Morotopithecus and elimination of distinct clades of
hominoids during this interval. The MMPC
accrues stratigraphic parsimony debt by implying a long missing lineage of hominoids in
a region that has been well sampled and from
which many hominoid localities are known. If
the morphological hypothesis is correct, then
the ‘‘modern’’ lineage is completely unrepresented in the fossil record, despite the presence of fossil localities in the geographic region and in strata of the correct age where
these hominoids should exist. If the stratocladistic hypothesis is correct, then the derived postcranial morphology that Morotopithecus shares with later hominoids of ‘‘modern
aspect’’ would be homoplasious. Although it
was demonstrated that the ‘‘archaic’’ clade
formed a monophyletic group unto itself even
upon the exclusion of Morotopithecus, the morphological support of a distinct clade of ‘‘archaic’’ hominoids is weak (low Bremer support and low bootstrap proportions). We favor
the stratocladistic result here, as it is unlikely
that an entire lineage of early Miocene hominoids is completely unrepresented in the relatively good fossil record of the early Miocene
of eastern Africa. New fossils from this region
will help to resolve this dispute not only because they would increase the chance of recovering such a lineage, but also because any
←
clades of ‘‘archaic’’ and ‘‘modern’’ hominoids. The previously recognized clade of ‘‘archaic’’ hominoids is now essentially grafted between the early appearance of Morotopithecus and the later FAE of Sivapithecus. D, Arrow points
to the elimination of stratigraphic debt by the displacement of Oreopithecus crownward. These differences in topology highlight areas of inconsistency between stratigraphic and morphological data. For instance, either there is
a lineage of ‘‘modern’’ hominoids unrepresented in the fossil record of eastern Africa, despite productive fossil
localities in the region, or the morphology observed in Morotopithecus linking this taxon to a clade of ‘‘modern’’
hominoids is homoplasious. Note also that the long ghost lineages between Hylobates and Pongo and their sister
groups persist in the stratocladistic hypothesis. These are not overturned by stratigraphic information because of
the strong morphological evidence for their respective positions.
634
JOHN A. FINARELLI AND WILLIAM C. CLYDE
new morphological information (especially in
the postcrania of such taxa as Morotopithecus
and Kenyapithecus) should bear heavily on
phylogenetic hypotheses.
The second interval of incongruence between morphology and stratigraphy occurs in
the late Miocene across Eurasia. For instance,
Oreopithecus is displaced crownward in the
cladogram, removing stratigraphic parsimony debt incurred by the MMPC. This late-appearing taxon is joined at the base of the hominid clade in the MMPC (Fig. 7A). If the morphological hypothesis is correct, then a large
sampling gap exists during this interval for
the lineage leading to Oreopithecus. There is
some evidence to believe that this may be the
case. Oreopithecus is associated with a fauna
that is a unique mixture of African and European taxa, and the localities from which this
taxon is known are thought to sample the remnant of a now-submerged land corridor between northern African and southern Europe
(Andrews et al. 1996; Harrison and Rook
1997). On the other hand, if the stratocladistic
result is correct, then Oreopithecus represents
the sister group to the Homininae. This implies a European origin for Oreopithecus (Andrews et al. 1996), and a southward migration
for the hominines into Africa, with Oreopithecus representing an island isolate that remained on the land bridge. The highly diverged morphology of Oreopithecus presents a
problem in the evaluation of this hypothesis.
Oreopithecus is placed at the base of all great
apes in the MMPC, sharing only primitive
characters with the remainder of the Hominidae, and bootstrap analyses could find no
strong support for resolving the polytomy at
the base of this clade. We suspect that the incongruence in this case results from the unusual stratigraphic setting of Oreopithecus,
which may greatly underestimate the FAE of
this lineage compared with coeval hominoid
taxa, in particular Sivapithecus, which likely
has a higher preservation potential given the
geologic setting in which the fossil material
has been recovered, and therefore likely has a
more accurate estimate of its FAE.
Conclusions
Several important conclusions can be drawn
from the phylogenetic analyses presented
here.
1. The morphological evidence points to the
presence of two distinct evolutionary lineages of hominoids in the earliest Miocene,
a postcranially primitive clade of ‘‘archaic’’
hominoids including early Miocene taxa
from eastern Africa, and a postcranially
derived clade of ‘‘modern’’ hominoids that
includes middle to late Miocene Eurasian
hominoids and all extant taxa. Morotopithecus is positioned at the base of the ‘‘modern’’ hominoid clade consistent with the
hypothesis of MacLatchy et al. (2000). However, the support for the separation of ‘‘archaic’’ hominoids from ‘‘modern’’ hominoids is shown to be rather weak by both
consensus and bootstrap methods. Evaluation of reduced consensus cladograms
and several bootstrapping analyses indicate that the cladistic hypothesis is very
sensitive to choice of ingroup taxa and that
the poor preservation of a few taxa (e.g.,
Kenyapithecus) is responsible for much of the
observed cladistic uncertainty.
2. The distribution of homoplasy across taxonomic and character partitions is highly
asymmetric. The ‘‘archaic’’ hominoid clade
displays very little homoplasy compared
with the ‘‘modern’’ hominoid clade. This
suggests that the elevated level of homoplasy observed in the Hominoidea is confined
largely to the clade of ‘‘modern’’ hominoids. In addition, the level of homoplasy
observed in dental characters relative to the
MMPC is significantly higher than that observed for postcranial characters. This result was upheld even when the analysis
was performed on a set of randomly generated rarefactions that were created to
equalize both preservation and implicit
weighting of the two character partitions.
Postcranial characters appear to be the
more reliable indicators of hominoid phylogeny under the parsimony criterion, indicating that they should become a focus of
future collection efforts.
3. A better-than-average match is observed
between the ordering of hominoid FAEs
and the branching order in the MMPC, indicating that the hominoid fossil record is
relatively good. Most of the observed mismatch is within the group of ‘‘modern’’
DATA CONGRUENCE IN HOMINOID PHYLOGENY
hominoids. However, when stratigraphy is
incorporated into the analysis there are significant topological differences between
the stratocladistic hypothesis and the
MMPC. These differences highlight two intervals in the stratigraphic record where
significant incongruence exists between the
morphological and stratigraphic data for
the Hominoidea. The first interval in the
early Miocene of eastern Africa is the result
of the morphological support for distinct
clades of ‘‘archaic’’ and ‘‘modern’’ hominoids. The early appearance of Morotopithecus implies an unobserved lineage of
‘‘modern’’ hominoids spanning an interval
that is otherwise characterized by good ingroup preservation. In light of the rather
weak morphological support for the position of Morotopithecus, the reality of two distinct evolutionary lineages in the early
Miocene is questionable. Rather it is likely
that the taxa in the ‘‘archaic’’ group here
represent a stem lineage of early hominoids. Additional material (especially postcrania) will certainly help to resolve the
evolutionary relationships of these early
hominoid taxa. The second interval in the
late Miocene of Eurasia is highlighted by
the variable position of Oreopithecus. In this
case, its morphology is well known but it
comes from a unique stratigraphic setting
in southern coastal Europe that makes the
temporal range of Oreopithecus and/or its
immediate ancestors poorly resolved compared to coeval hominoid taxa.
Acknowledgments
We would like to thank W. Sanders for helpful insights in the course of our analysis, and
D. Fisher and D. Begun for thoughtful and
critical reviews of our manuscript, which have
greatly improved the quality of this work. In
addition, the work presented here would not
have been possible without the support of R.
Potts and J. Clark at the Human Origins Program, Smithsonian Institution and A. Rosenberger then at the Smithsonian Institution.
Partial funding for this work was provided by
the University of New Hampshire Graduate
Teaching Assistant Summer Fellowship and
the Jonathon W. Herndon Scholarship in Earth
635
Sciences. This research was part of a Master’s
Thesis that J.A.F. submitted to the Graduate
School at the University of New Hampshire.
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ovoid
broad
angular
retroflexed
subequal
distinct
straight
small
separate
small
lateral
fused
large
palmar
rounded
present
circular
concave
laterally
reduced
broad
shallow
abbreviated
spherical
narrow
rounded
medial
.medially
merged
notched
large
,90 deg.
,15 deg.
oval
absent
oval
flat
anterolaterally
prominent
narrow
deep
not abbreviated
.90 deg.
.15 deg.
retroflexed
Anteroposterior shaft
curvature
Intertuberosity angle
Humeral head torsion
Humeral head shape
Bicipital groove
Proximal shaft shape
Medial epicondylar projection
Trochlear keel symmetry
Medial epicondyle/keel
Superior trochlear border
Lateral trochlear keel
Radial head shape
Radial head beveling
Radial neck shape
Distal radial articular surface
Radial notch of ulna faces
Deltopectoral plane
Trochlear notch
Ulnar shaft
Olecranon process
larger
deep
trochleiform
w/strong
medial keel
straight
narrow
anterior
gracile
smaller
1
smaller
shallow
cylindrical w/
weak medial keel
broad
axillary
robust
larger
0
Character state definitions
Appendix 1
Coronoid/radial fossae size
Coronoid fossa
Trochlea
Infraglenoid tubercle
Teres minor attachment
Spinous process root
Glenoid/axillary margin
angle
Carpo-metacarpals
28
Os centrale and scaphoid
29
Scaphoid tuberosity
30
Scaphoid tubercle
Radius/Ulna
19
20
21
22
23
24
25
26
27
9
10
11
12
13
14
15
16
17
18
8
Humerus
5
6
7
Scapula
1
2
3
4
Character number and description
Morphological character descriptions.
2
3
et
et
et
et
al.
al.
al.
al.
1997a
1997a
1997a
1997a
et
et
et
et
et
et
et
et
et
et
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al.
Begun et al.
Begun et al.
Begun et al.
Rose 1997
Begun et al.
Begun et al.
Begun et al.
Rose 1997
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun
Begun
Begun
Begun
Original
citation
34
37
38
30
31
32
33
z/z9
168
170
171
y/y9
21
22
23
24
25
26
27
28
29
169
20
16
17
18
2
3
4
5
Number in
original
citation
DATA CONGRUENCE IN HOMINOID PHYLOGENY
639
Os centrale articulation with
trapezium
Lunate: mediolateral width
Lunate: dorsopalmar length
Scaphoid facet on lunate
Lunate, scaphoid radial facet
Ulnar styloid articulation with
pisiform and/or triquetrum
MC3 facet on capitate
Palmar MC4 facets on capitate
MC2 facet on capitate
Trapezoid facet on capitate
Capitate head mediolaterally
Central facet on capitate
Hamulus size
Triquetral face on hamate
MC4 facet on hamate
MC1 head proximodistally
MC1 head dorsal part
MC1/trapezium articular
surface
MC4 palmar capitate facet
MC facets on palmar hamate
MC2-4 sesamoids
MC heads broadest
Trunk
56
57
58
59
60
61
Costal angle
Vertebral body height
Accessory processes
Transverse processes
Sternebrae
Torso shape
Phalanges: Manus
53
Proximal articulation surface
of proximal phalanges
Proximal phalangeal palmar
54
tubercles
Ray 1 terminal phalanx
55
articulation surface
49
50
51
52
37
38
39
40
41
42
43
44
45
46
47
48
32
33
34
35
36
31
Continued.
Character number and description
Appendix 1.
ridged
convex
high
medium
small
inter.
inter.
short/mediolaterally
broad
small
large
low
tall
large
ventral
narrow
craniocaudally
long/narrow
square
absent
angulated
absent
dorsally
oval
present
confluent
present
palmarly
et
et
et
et
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Ward, C.V. 1997
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun
Begun
Begun
Begun
et
et
et
et
et
et
et
et
et
et
et
et
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
irregular
absent
divided
present
narrow
concavoconvex
large
lateral
shallow
short
bowed
proximal
flat
present
continuous
absent
broad
flat
small
proximal
deep
long
narrow
dorsal
al.
al.
al.
al.
al.
Begun et al. 1997a
et
et
et
et
et
3
Begun
Begun
Begun
Begun
Begun
short
absent
dorsal
broad
2
broad
short
extensive
high-angle
absent
absent
1
Original
citation
narrow
long
restricted
low-angle
present
present
0
Character state definitions
74
75
76
77
79
In text
73
72
71
65
66
67
68
52
53
54
55
56
57
58
59
60
61
62
63
42
43
44
45
47
39
Number in
original
citation
640
JOHN A. FINARELLI AND WILLIAM C. CLYDE
Distal tibia facet
Medial malleolus projection
Fibular robusticity
Lateral malleolus
Tibia/Fibula
71
72
73
74
Tarso-metatarsals
75
Talar trochlear depth
76
Distal calcaneus
77
Flexor hallucis longus groove
78
Posterior talar facet long axis
79
Anterior talar facet
80
Plantar calcaneal tubercle
Calcaneo-navicular facet
81
Cuboid peg
82
Cuboid wedging
83
Cuboid length
84
Entocuneiform/MT 1 joint
85
Cuneiform length
86
MT 1 robusticity
87
MT 1 sesamoid grooves
88
MT 1 length
89
MT 1 prehallux facet
90
MT 2-5 robusticity
91
Transverse arch
92
Entocuneiform facet on
93
navicular
Trochanteric fossa
Femoral head
Femoral neck tubercle
Femoral condyle shape
Iliac blade width
Iliac blade height
Lower iliac height
Cranial aspect of acetabulum
lunate surface
Pubic length
Femur
67
68
69
70
66
Pelvis
62
63
64
65
Continued.
Character number and description
Appendix 1.
short
deep
long
small
aligned
curved
small
large
small
slight
long
distal
long
gracile
small
long
present
gracile
present
dorsal
square
distal
thin
small
open
cylinder
present
symmetrical
narrow
low
short
narrow
0
inter.
inter.
inter.
angled
inter.
large
small
inter.
strong
short
medial
short
inter.
large
short
absent
robust
absent
inter.
short
flared
robust
inter.
distal
robust
large
flat
shallow
short
large
large
asymmetrical
et
et
et
et
et
et
et
et
al.
al.
al.
al.
al.
al.
al.
al.
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
Ward, C. V. 1997
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
94
98
99
101
102
103
104
105
106
107
108
109
110
111
114
115
116
117
41
90
91
92
93
85
86
87
89
long
medium
deep
sphere
84
Begun et al. 1997a
wide
inter.
inter.
absent
inter.
80
81
82
83
al.
al.
al.
al.
1997a
1997a
1997a
1997a
et
et
et
et
Begun
Begun
Begun
Begun
unique
3
Number in
original
citation
wide
2
Original
citation
inter.
high
long
inter.
1
Character state definitions
DATA CONGRUENCE IN HOMINOID PHYLOGENY
641
Continued.
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
114
Lower central incisors
Lower incisors labiolingually
Lower canine crown height
Lower molar cingula
M1/M2 size ratio
M3/M2 size ratio
I1/I2 size heteromorphy
Upper canine height
Upper canine cervical flare
P4 shape
P3 shape
Premolar buccal flare
P4 lingual flare
M1–2 shape
Upper molar cingula
buccolingually
oriented
large
small
tall
strong
small
M3.M2
low
tall
weak
broad
triangle
weak
weak
broad
strong
broad
present
large
robust
narrow
strong
tall/narrow
narrow
absent
absent
short
low
large
tall
high
subequal
Dentition
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
Incisor breadth
I2 cingulum
Relative male canine size
Male canines
Canine cingula
P3 cusp heteromorphy
P3 paracones
P3 shape
P3 mesiolingual beak
P3 metaconid
P4 shape
P4 talonid
M2–3 metacones
Molar cusps
Dentine penetrance
Anterior dentition relative to
posterior dentition
Mandibular canine roots
digit 3
gracile
straight
weak
0
Phalanges: Pes
94
Foot axis runs through
95
Phalangeal robusticity
96
Phalangeal curvature
97
Phalangeal flexor ridges
Character number and description
Appendix 1.
narrow
absent
reduced
compressed
thick, rounded
reduced
low/rounded
broad
present
present
long
high
reduced
low/rounded
low/rounded
larger postcanine
internally
rotated
small
large
short
weak/absent
large
M3,/5M2
high
low/rounded
strong
long
rectangle
strong
strong
long
weak/absent
digit 2
robust
curved
inter.
1
2
v. high
v. low
v. reduced
strong
Character state definitions
3
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
218
219
220
221
222
223
224
226
227
228
229
230
231
232
234
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
118
119
120
121
217
1997a
1997a
1997a
1997a
Begun et al. 1997a
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
Number in
original
citation
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Original
citation
642
JOHN A. FINARELLI AND WILLIAM C. CLYDE
161
162
163
160
159
154
155
156
157
158
153
148
149
150
151
152
Continued.
External occipital
protuberance
Nuchal plane orientation
Neurocranial length
Glenoid fossa
Articular tubercle
Articular/tympanic and
temporal bones
Zygomatic root pneumatization
Inferior orbital foramen
relative to nasal aperture
Zygomatic depth
Zygomatic orientation
Maxillary nasal process
Palatine process
Alveolar process
Incisive fossa
Maxillary depth
Lateral malar surface
Frontozygomatic breadth
Orbital breadth
Nasal bones at nasion
Glabella
Supraorbital torus
Supraciliary ridges
Supraorbital sulcus
Frontal sinus size relative to
nasion
Frontal sinus size
Frontal squama
Facial profile
Temporal fossa
Inion relative to glabella
M1/M2 size ratio
Upper molar crowns
Upper molar sides
Molar enamel
M3 size
Character number and description
Cranium
135
136
137
138
139
140
141
142
143
144
145
146
147
130
131
132
133
134
Appendix 1.
1
shallow
lateral
robust
deep
anterior
hollow
al.
al.
al.
al.
al.
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
et
et
et
et
et
178
175
177
173
Begun
Begun
Begun
Begun
Begun
Begun et al. 1997a
al.
al.
al.
al.
al.
below apex
near apex
et
et
et
et
et
172
Begun
Begun
Begun
Begun
Begun
Begun et al. 1997a
solid
inferior
138
139
140
144
145
146
147
148
149
150
151
152
153
235
236
237
238
239
159
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun et al. 1997a
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
154
155
156
157
158
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
1997a
1997a
1997a
1997a
1997a
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
hollow
posterior
elongated
deep
small
fused
superiorly broad
short
shallow
large
unfused
v. large
3
Number in
original
citation
160
162
163
164
167
reduced
strong
inion significantly lower
absent
absent
sunken
strong
v. elongate
distal to P3
2
Original
citation
1997a
1997a
1997a
1997a
1997a
small
horizontal
concave
broad
inion below
large
vertical
convex/flat
narrow
inion above
large
high
vertical
crenulated
reduced
thick
biconvex
distal to C
deep
flat
thicker
elongate
narrow
inflated
present
present
broad
above & below
0
thin
flat
opposite to C
shallow
curved
narrow
broad
broad
indistinct
absent
absent
shallow/absent
above
small
low
bulging
smooth
large
Character state definitions
DATA CONGRUENCE IN HOMINOID PHYLOGENY
643
Continued.
Maxillary alveolar process
Incisor orientation
Greater palatine foramen
Greater palatine position
Lesser palatine foramina
Horizontal palatine
Palatine crest
Posterior palate
Pyramidal process position
Pterygoid process
Alveolar process depth
Zygomatico-alveolar crest
Zygomatic root height
Incisive fossa
Subnasal floor
Nasoalveolar clivus length
Nasoalveolar clivus
orientation
Incisive canal caliber
Nasal aperture shape
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
177
178
179
180
181
176
173
174
175
172
165
166
167
168
169
170
171
Orbital/nasal aperture
distance
Orbital/nasal surface
Interorbital distance
Zygomatic arch relative orbit
Zygomatic temporal process
Zygomatic arch angle
Nasal aperture breadth
Nasal aperture relative to orbit
position
Nasal aperture relative to
alveolar plane
Lacrimal fossa visible
Nasal bone length
Nasal aperture relative to
malar surface
Inferior orbital margin relative
to M1
Nasal apex relative to M2
Maxillary surface
Canine fossa
Canine foot angulation
Canine root rotation
164
Character number and description
Appendix 1.
large
ovid
solid
vertical
round
anterior
none
broad
strong
shallow
superiorly broad
robust
shallow
compressed
low
absent
fenestrated
short
vertical
absent
superiorly broad
posterior
posterior
anterior
deeper
medial
externally
rotated
collapsed
horizontal
elongate
posterior
small
narrow
weak
deep
inferior
compressed
deep
broad
high
shallow
stepped
long
horizontal
above
lateral
shallow
vertical
in line
above
small
piriform
deep
smooth
large
inflated
v. long
flat
v. small
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
al.
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
1997a
Begun et al. 1997a
Begun and Güleç 1998
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun et al. 1997a
216
18
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
214
215
193
194
195
196
197
192
189
190
191
Begun et al. 1997a
Begun et al. 1997a
Begun et al. 1997a
yes
long
inter.
none
short
anterior
al.
al.
al.
al.
al.
al.
al.
187
et
et
et
et
et
et
et
178
Begun et al. 1997a
Begun
Begun
Begun
Begun
Begun
Begun
Begun
Begun et al. 1997a
high
low
high
horizontal
3
Number in
original
citation
179
180
182
183
184
185
186
v. large
2
Original
citation
1997a
1997a
1997a
1997a
1997a
1997a
1997a
flat
narrow
below
deep
vertical
broad
v. low
small
1
concave
broad
same level
shallow
inclined
narrow
low
large
0
Character state definitions
644
JOHN A. FINARELLI AND WILLIAM C. CLYDE
1 1 1
0 1 1
0 0 0
0 0 0
character
Hylobates
1 1 0 1
1 1 1 1
1 0 0 0
0 0 1 0
Stratigraphic
1 1
1 1
0 0
1 0
‘‘k’’
1 1
2 1
1 1
1 0
‘‘k’’
1 1
2 1
1 0
0 0
‘‘k’’
1 1
2 1
1 0
1 0
‘‘k’’
1 1
2 1
1 1
1 0
‘‘j&k’’
1
1
0
0
1
2
0
1
1
2
0
0
1
2
0
0
1
2
1
1
1
1
0
0
1
1
0
1
1
1
0
1
1
1
0
1
1
1
1
1
0 — 0 — 0 0
1 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
state: ‘‘a&b&c&d’’
1 1
1 1
0 0
0 0
state:
1 1
2 2
1 1
0 1
state:
1 1
2 2
1 1
1 1
state:
1 1 1
1 1 2
1 1 1
0 0 0
character
Proconsul
0 0 1 0 0 0 0
0 0 0 0 — 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
Stratigraphic character
Character number
Appendix 2
0
0
0
0
1
1
0
1
1
1
1
1
1
1
0
0
1
1
0
1
1
0
0
1
0
0
0
0
0
1
0
1
1
2
0
1
1
2
0
0
1
2
0
0
1
2
0
0
1
0
0
0
0
1
0
0
0
2
1
1
0
2
1
0
0
2
1
0
0
2
1
0
0
0
0
0
0
1
0
1
1
2
0
1
1
2
0
1
1
2
0
1
1
3
2
0
0
1
0
0
1
2
1
0
1
2
1
0
1
2
1
0
1
2
1
0
1
2
1
0
0
0
0
0
1
0
0
0
1
1
1
1
1
1
1
2
1
1
1
2
1
1
1
2
1
1
0
0
1
0
1
0
1
2
1
1
1
2
1
1
1
2
1
0
1
2
1
0
1
0
0
0
1
1
0
2
1
1
2
2
1
1
0
0
1
1
0
2
1
0
1
2
0
0
0
0
1
1
0
0
1
1
0
1
1
1
0
1
1
1
0
1
1
1
2
1
0
1
0
0
1
0
0
0
1
1
0
0
1
1
0
1
1
1
0
1
1
1
0
1
0
1
0
0
1
2
0
0
1
2
1
1
1
2
1
1
1
2
1
0
1
0
1
0
0
1
0
0
1
2
0
0
1
2
0
2
1
2
0
1
1
2
0
1
1
2
1
1
0
0
0
0
1
1
0
0
1
2
0
1
1
2
0
0
1
2
0
0
1
2
0
0
0
1
0
0
1
1
0
0
1
2
0
1
1
2
0
0
1
2
1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
0
1
1
1
1
1
1
1
1
0
1
0
0
1
2
1
1
0
2
1
1
1
2
1
0
1
2
1
0
1
2
1
1
0
0
0
0
1
0
1
0
1
1
1
1
1
1
1
0
1
1
1
0
1
1
1
0
0
0
0
0
1
1
0
0
1
1
1
0
1
1
1
0
1
1
1
0
1
1
1
1
0
2
0
0
1
1
0
0
1
2
1
1
1
1
1
2
1
1
1
2
1
1
1
2
0
0
0
0
0
0
0
0
1
1
1
1
1
1
0
0
1
1
0
0
1
1
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
0
0
1
1
1
0
1
0
0
0
0
1
0
0
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
0
0
1
0
0
0
1
1
1
1
1
1
1
2
1
0
1
2
0
1
0
0
0
1
0
0
1
2
1
1
1
2
1
1
1
2
2
1
1
2
2
1
0
1
0
0
1
0
0
0
1
0
1
0
1
0
1
1
1
0
1
1
1
1
1
0
0
0
0
0
1
1
0
1
1
1
1
1
1
1
0
1
1
1
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
1
1
1
0
1
1
1
0
1
0
1
0
1
0
1
0
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
0
0
1
1
1
1
1
1
1
1
1
2
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1 0 1
0 — —
0 0 0
0 0 0
1
1
2
0
0
1
0
1
0
1
1
1
0
1
1
1
1
0
0
0
1
0
0
0
1
1
0
1
0
1
2
1
1
1
2
1
1
1
2
1
0
0
0
0
1
0
0
0
1
1
1
1
0
1
1
2
1
1
1
2
1
1
1
2
0
0
0
0
1
1
0
0
0
1
0
2
1
1
1
1
1
1
1
1
1
0
1
1
0
1
0
0
1
2
1
0
1
2
2
2
1
2
1
1
1
2
1
2
1
2
1
2
0
0
0
0
1
0
0
0
1
0
2
1
1
0
3
0
1
0
3
0
1
0
3
0
0
0
0
0
1
0
0
0
1
1
0
3
1
1
1
2
1
1
1
2
1
1
0
2
0
0
0
0
1
0
0
0
1
1
1
2
1
1
0
1
1
1
0
1
1
2
0
1
9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Pongo
1 1 0 1
1 1 1 1
1 1 1 1
0 1 1 1
Stratigraphic
8
1 1 1
1 1 2
1 1 1
1 1 1
character
7
Gorilla
1 1 0 1
1 1 1 1
1 1 1 1
1 1 1 2
Stratigraphic
6
1 1
2 2
1 1
1 1
state:
5
1 1 1
1 1 2
1 1 1
1 1 1
character
4
Pan
1 1 0 1
1 1 1 1
1 1 1 1
1 2 1 2
Stratigraphic
3
1 1
2 2
1 1
1 1
state:
2
Australopithecus
1 1 0 1 1 1 1
1 0 1 1 1 1 2
1 1 1 1 1 1 1
1 2 1 2 1 1 1
Stratigraphic character
1
Character by taxon matrix for the Hominoidea.
DATA CONGRUENCE IN HOMINOID PHYLOGENY
645
Character number
0
—
0
—
1
—
0
0
Griphopithecus
— — — — — — 0 0 — — — —
— — — — — — — — — — — —
0 1 1 1 1 1 1 1 1 1 1 1
— — — — — — — — — — — —
Stratigraphic character state: ‘‘c&d’’
Dryopithecus
— — — — 1 1 1 1 — — —
— — — — — — — — 2 — —
1 1 1 1 1 1 1 1 1 1 0
1 1 1 1 1 1 1 1 1 1 —
Stratigraphic character state: ‘‘f&g’’
—
—
0
0
0
—
0
—
0 0
1 0
0 1
— —
‘‘c’’
—
—
0
—
—
—
0
—
—
—
0
—
1
—
0
—
—
—
—
—
0
—
0
—
0
—
—
—
—
—
0
2
—
—
—
2
1
—
0
0
—
—
1
—
0
—
0
—
0
—
0
—
—
—
0
0
0
—
—
0
0
—
1
0
—
—
1
—
—
—
—
—
0
—
—
—
—
—
0
0
—
—
—
0
1
—
0
—
0
—
1
—
—
0
1
—
1
—
0
—
—
—
1
0
—
0
—
1
1
—
1
0
0
—
0
—
0
1
1
—
0
—
—
—
0
—
0
—
—
1
—
0
1
—
1
0
—
—
0
—
1
0
0
—
—
—
—
—
—
—
0
1
1
—
—
1
1
—
1
0
—
—
1
—
—
—
0
—
—
—
—
—
—
—
0
0
—
1
1
—
—
—
1
—
—
0
1
—
—
—
1
—
—
—
0
1
—
0
0
—
—
—
0
—
0
0
—
—
—
—
0
—
—
—
0
0
1
—
0
1
1
—
0
—
1
1
—
—
—
—
0
—
—
1
0
1
0 0 0 0
1 — — —
0 — 0 —
0 1 0 —
1
—
0
—
0
—
0
—
0
1
0
—
—
—
0
—
0
1
0
1
—
—
0
1
0
—
0
0
0
—
—
0
1
1
0
—
0
—
0
—
0
1
0
—
—
—
0
—
1
—
0
—
0
—
0
—
0
—
—
—
—
—
0
—
—
1
0
—
0
—
0
—
0
1
—
—
—
—
0
—
0 0 —
1 — —
0 1 0
0 0 0
—
1
0
0
—
—
1
0
—
—
1
—
0
0
0
—
—
—
0
—
0
—
0
0
—
—
0
0
—
—
1
0
—
—
1
1
—
1
1
—
—
—
1
1
—
—
0
1
—
—
1
1
—
—
1
0
—
—
0
—
—
0
0
—
—
—
0
1
—
—
0
0
—
—
0
0
—
—
1
0
—
—
0
—
—
0
0
—
—
—
0
1
—
—
0
0
—
—
—
0
1
—
1
0
—
—
1
1
—
—
1
1
—
—
1
1
0
2
0
0
—
2
1
0
—
0
1
—
1
—
0
0
—
—
—
—
—
—
0
0
—
—
0
—
1
—
0
—
—
—
1
—
—
—
0
—
—
—
—
—
0 0
0 0
0 1
0 —
—
0
0
0
0
1
1
—
—
—
1
—
—
0
—
—
—
—
1
—
0
—
—
0
—
—
—
—
—
0
1
—
—
—
—
—
0
—
—
—
—
—
—
—
—
—
0
0
0
—
—
0
—
—
1
—
—
—
—
—
—
1
—
—
—
—
0
—
0
—
0
0
—
—
—
—
—
—
0
—
—
—
0
—
—
—
1
—
—
—
0
—
—
1
0
—
—
—
0
—
—
—
0
1
—
—
—
—
—
0
—
—
—
—
—
—
—
—
1
—
—
—
—
—
—
1
—
—
—
—
—
—
0 — —
1 0 0
1 0 0
0 0 0
—
—
0
—
—
1
1
—
—
—
—
—
—
0
—
—
—
—
—
—
0
0
1
0
—
—
0
—
—
—
—
0
—
—
1
0
—
—
0
1
—
—
1
1
—
—
—
0
—
—
—
1
—
—
—
1
—
—
—
0
1
0
—
0
—
—
—
1
—
—
—
—
0
0
—
—
—
—
—
—
—
—
—
—
—
0
—
—
—
—
—
—
—
0
0
—
—
—
0
—
—
—
—
—
—
1
—
—
—
—
—
—
—
1
0
1
—
—
0
1
1 1 1 — —
0 — 1 1 1
1 1 1 0 1
1 1 2 1 1
—
—
—
—
—
—
—
—
—
—
—
—
0 — — —
0 — 0 0
0 0 0 0
0 0 0 —
—
—
0
0
—
1
1
0
—
0
—
—
—
0
—
—
—
0
—
—
—
0
0
0
—
—
0
0
—
1
0
1
—
0
—
—
—
0
—
—
—
0
—
—
0
0
0
—
—
—
0
0
—
1
0
—
—
1
—
—
—
0
—
—
—
1
—
—
0
0
0
2
—
—
0
0
9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
0
—
1
1
8
Equatorius
— — — — — — — 0 0
1 0 0 0 0 1 0 — 0
0 0 1 1 1 1 1 1 1
— — — — — — — — 1
Stratigraphic character state:
7
—
—
1
1
6
Kenyapithecus
— — — — 0 0 0 — — — —
— — — — — — — — — — —
1 1 1 1 1 1 1 1 1 0 1
— — — — — — — — — — —
Stratigraphic character state: ‘‘d’’
5
—
—
—
0
4
Afropithecus
— — — — — — — — — — —
0 — 0 0 0 — — — — — —
0 0 0 0 1 0 0 0 — 0 1
— — — — — — — — — 0 1
Stratigraphic character state: ‘‘b’’
3
—
—
—
0
2
Continued.
Turkanapithecus
— — — — — — 0 — — — —
— — 0 — — — — — — — 0
— — — — — — — — — — —
1 — 0 — — 0 — — — 0 1
Stratigraphic character state: ‘‘b’’
1
Appendix 2.
646
JOHN A. FINARELLI AND WILLIAM C. CLYDE
2
3
4
5
6
7
8
Continued.
1 1
2 2
1 0
1 1
state:
1 1
2 1
1 0
1 —
‘‘i’’
—
—
1
—
—
—
0
—
Ouranopithecus
— — — — — — — — — — — —
— — — — — — — — — — — —
1 1 1 1 1 1 1 1 1 1 1 1
— — — — — — — — — 1 — 0
Stratigraphic character state: ‘‘g&h’’
—
—
—
—
—
—
1
1
Morotopithecus
— — — — — — — — — — —
— — — — — — 2 2 2 — —
— 0 — — — — — — — 0 —
— — — — — — — — 1 — —
Stratigraphic character state: ‘‘a’’
Ankarapithecus
— — — — — — — — — — —
— — — — — — — — — — —
0 1 1 1 0 1 1 0 0 1 1
— — — — — — — — 0 0 1
Stratigraphic character state: ‘‘f’’
0
—
1
1
1
1
0
0
1 — —
1 — 2
1 0 1
0 1 0
1
—
2
1
—
0
0
1
—
—
0
0
—
—
—
—
—
—
0
—
—
—
0
0
—
—
1
0
—
—
—
—
—
—
0
0
—
—
0
0
—
—
1
1
—
—
—
0
—
—
1
0
—
—
1
0
—
—
1
—
—
0
—
—
—
—
2
—
—
—
0
0
—
—
1
1
—
1
—
—
—
—
1
0
—
—
1
0
—
—
0
1
—
0
—
—
—
—
1
0
1
—
1
1
—
—
1
1
—
2
—
0
—
—
0
0
—
—
1
0
—
—
2
1
—
—
0
0
—
—
1
—
—
—
2
2
—
—
1
1
—
—
0
0
—
—
1
1
—
—
0
0
1 1 0 — 1 1 1 — —
1 — 2 1 2 — 2 — 1
0 0 0 0 1 1 0 1 0
1 0 0 0 1 1 0 2 1
0
—
0
1
0
—
0
2
—
—
1
1
—
—
0
—
—
—
1
1
—
—
0
1
—
—
0
2
—
—
0
—
—
—
0
1
—
—
0
0
—
—
0
0
—
—
0
0
—
—
1
1
—
—
1
1
—
—
0
0
—
—
1
0
—
—
0
0
—
—
0
0
1
2
0
0
—
0
1
1
—
2
1
1
—
—
0
0
—
—
0
0
—
—
1
—
—
—
0
0
—
—
1
0
—
—
0
0
—
—
1
0
—
—
1
0
—
—
1
1
—
—
0
0
—
—
1
0
—
—
1
1
1 — —
2 0 2
0 1 0
0 0 0
1 — —
1 1 2
0 0 0
2 1 0
1 1 1
1 — 2
0 0 1
0 0 —
—
—
0
0
—
—
1
1
—
—
0
0
—
—
1
1
—
—
1
1
—
1
1
0
—
1
1
1
—
—
1
1
—
—
0
0
—
—
1
1
—
—
1
—
—
1
1
0
—
1
1
1
—
—
1
0
—
—
1
0
—
—
1
1
—
—
1
0
—
2
1
0
—
2
1
1
—
—
0
0
—
—
—
0
—
—
0
0
—
—
1
0
—
1
0
0
—
0
0
1
—
—
0
1
—
—
—
—
—
—
0
—
—
—
—
—
—
1
0
—
—
1
0
1
—
—
1
1
—
—
1
0
—
—
1
—
—
—
—
—
—
1
—
—
—
—
1
1
—
—
1
0
—
—
0
0
—
—
1
—
—
—
1
—
1
—
—
—
—
0
0
—
—
—
0
1
—
—
0
0
—
—
1
—
—
—
—
—
—
—
—
—
—
2
1
1
—
—
1
0
—
—
0
0
—
—
1
—
—
—
—
—
—
—
—
—
—
0
1
0
—
—
—
1
—
—
—
0
—
—
0
1
—
—
0
—
—
0
0
—
—
0
1
1
—
—
—
0
—
—
—
0
—
—
1
—
—
—
1
—
—
1
0
—
—
1
0
0
—
—
0
0
—
—
—
0
—
—
1
—
—
—
0
–
–
1
0
—
—
1
2
1
—
—
1
1
—
—
—
0
—
—
1
1
—
—
0
1
—
1
—
0
—
0
1
1
—
—
0
0
—
—
—
1
—
—
1
1
—
—
2
—
1
0
1
1
—
—
1
1
—
—
—
0
—
—
1
0
—
—
0
1
—
—
—
2
—
—
—
0
—
1
1
2
—
—
0
—
—
—
0
1
—
—
—
0
—
1
0
1
—
—
0
—
—
—
1
2
—
—
—
0
—
1
—
1
—
—
—
1
—
0
3
1
—
—
—
0
—
0
—
0
—
1
—
0
—
1
0
2
—
—
—
0
—
1
0
1
—
0
0
—
—
0
0
1
—
1
0
2
—
—
—
—
—
2
0
1
—
1
0
2
—
1
0
0
1 — —
0 1 1
2 0 1
1 3 2
1 — — — —
1 1 1 1 0
1 1 0 — 1
0 2 — 1 0
0 0 0 — —
1 — 1 1 1
0 0 1 0 2
0 1 1 2 2
* Characters in this table are arranged in four rows of fifty characters each (sum 200 characters). The first row contains 1–50; the second, 51–100; the third, 101–150 and the fourth, 151–200. The number in a particular
cell position gives the coded value of the character for the particular taxon. A dash denotes missing data.
—
—
1
0
—
—
0
1
1 1
2 1
0 0
0 —
—
—
—
1
Lufengpithecus
— — — 0 — — — — — — —
— — 0 0 — — — — — — —
1 — 1 1 1 1 1 1 1 — 1
— — — — — — — — — 1 0
Stratigraphic character state: ‘‘i’’
Oreopithecus
1 1 0 1 — — 1
— — — — — 1 2
1 — 1 1 1 0 1
— — — — 1 0 1
Stratigraphic character
Character number
9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Sivapithecus
— — — — 1 1 1 0 — 0 — 0 0 0
— — 0 0 — — — — — — 0 — — —
1 1 1 1 1 1 1 1 1 1 1 1 0 1
0 — — — — 0 0 0 0 0 1 1 1 0
Stratigraphic character state: ‘‘e&f&g&h&i’’
1
Appendix 2.
DATA CONGRUENCE IN HOMINOID PHYLOGENY
647
648
JOHN A. FINARELLI AND WILLIAM C. CLYDE
Appendix 3
Chronostratigraphy of the Hominoidea
Morotopithecus
Until recently, the age of the Moroto localities was thought to
be significantly younger than either of the two genera with
which the hominoid material was originally allied in the literature: Proconsul and Afropithecus. Bishop et al. (1969) reported
a K/Ar age for the beds at Moroto of 14.3 6 0.3 Ma. Pickford et
al. (1986) had revised this date on the basis of biochronological
correlations to older than 17.5 Ma, and recent revision of the
radiometric dates using the more precise 40Ar/39Ar method
places the age of the Moroto I and II localities at older than 20.61
6 0.05 Ma (Gebo et al. 1997).
Proconsul
Dating of Proconsul fossil material is generally well constrained as a result of the geological setting in East Africa ca.
20 Ma. The beginnings of the East African Rift resulted in the
formation of large volcanoes that produced tuffs intercalated
with the fossiliferous sedimentary strata, providing radiometric
dates bracketing hominoid-bearing units.
The fossil record of Proconsul in Kenya is known from the Tinderet and Kisingiri. Both have been reconstructed as sedimentary basins on the flanks of large shield volcanoes (Pickford and
Andrews 1981; Retallack 1991; Retallack et al. 1995). The Tinderet includes multiple fossil localities, including Koru, Leget,
Songhor, and Chamtwara. The stratigraphic sequence of interest includes the Koru Formation, overlain by the Leget Formation, and no fewer than 23 fossil sites bearing hominoid material
have been reported in these formations (Pickford 1984). This sequence is bracketed by unconformities, and K/Ar dating places
these beds between 19.5 and 19.6 Ma (Pickford and Andrews
1981). Above these beds lie the Kapurtay Agglomerates, including Songhor and Chamtwara. The Chamtwara Member has been
dated to ca. 19.7 Ma (K/Ar), and Songhor correlates stratigraphically to just above the Leget Formation (Pickford and Andrews
1981). Although a slight discrepancy appears to exist in the
dates, it is within the margin of error and supports a 19.7–19.5
Ma date for the Tinderet.
The Kisingiri localities consist of fossil sites on Rusinga and
Mfangano Islands in Lake Victoria. At least eight fossil sites
have been reported on Rusinga Island, including one site that
has yielded nine individuals of the species P. nyanzae (Walker
and Teaford 1988; Walker et al. 1993). Proconsul is known from
the uppermost portion of the Kinhera Formation, the entire Rusinga Agglomerate, and the earliest portion of the Hiwegi Formation, which form a conformable sequence of strata (Retallack
et al. 1995). Drake et al. (1988) reported K/Ar dates for tuffs in
the Rusinga Agglomerates and the Hiwegi Formations of 17.9 6
0.1 and 17.8 6 0.2 Ma respectively. Three sites on Mfangano Island have yielded hominoid fossils (Ward et al. 1993) and correlate to the Hiwegi Formation on Rusinga Island (Drake et al.
1988). Proconsul material has been recovered from the Napak
Formation in Uganda (Allbrook and Bishop 1963). This locality
was dated to between 18 and 19 Ma (Bishop et al. 1969).
Outside of the well-established localities in the Tinderet and
Kisingiri, Andrews et al. (1981) reported fossils that they assigned to Proconsul from the latest Oligocene/earliest Miocene
Muhoroni Agglomerates at Meswa Bridge, Kenya. These deposits have been dated to 23.5 Ma (Pickford and Andrews 1981; Tassy and Pickford 1983). In addition, several sites in the Ngorora
Formation of the Tugen Hills have produced fragmentary fossil
remains of primates that have been tentatively assigned to the
genus Proconsul. These beds in the Ngorora Formation have
been dated through a variety of techniques (K/Ar, 40Ar/39Ar,
paleobotanical correlation) to approximately 12.5 Ma (Deino et
al. 1990; Hill et al. 1985, 2002). Using the most encompassing
dates for Proconsul then places the FAE at Meswa Bridge, and
the LAE in the Ngorora Formation. However, although the taxonomic assignment of the Kisingiri and Tinderet material is
rather secure, the attribution of the hominoid fossils from the
Ngorora Formation and Meswa Bridge to Proconsul is less certain (see Andrews et al. 1981; Senut et al. 2000; Hill et al. 2002).
Thus, both the FAE and LAE of the OTU Proconsul, as used here,
may change dramatically with the recovery of more complete
material. The effect of these changes on the results of this analysis is minimal, however, and the topology of the stratocladistic
hypothesis is not affected. See text for discussion.
An additional locality exists that is purported to extend the
range of Proconsul into the latest Oligocene. Material from the
Eragaleit Beds of the Lothidok Formation was also referred to
Proconsul in beds dated to 24.3–27.5 Ma (Brochetto et al. 1992).
This material has been reassigned to the genus Kamoyapithecus,
a late Oligocene catarrhine (Leakey et al. 1995b). Although the
authors did note morphological affinities of this taxon to the genus Proconsul and tentatively placed this taxon within the Hominoidea, they pointed out that the level of preservation for this
taxon is as yet insufficient for use in cladistic analysis (Leakey
et al. 1995b), and consequently Kamoyapithecus is not considered
in the present study.
Afropithecus
Afropithecus is known from the region around Lake Turkana
from the early Miocene. The genus was described from material
recovered at the Kalodirr locality in Kenya (Leakey and Leakey
1986a) consisting of the single species Afropithecus turkanensis.
Material recovered from the Warata Formation at Buluk (Leakey
and Walker 1985) and Locherangan, Kenya (Anyonge 1991), is
also referred to this genus. Fossils from Morourot Hill (Andrews 1978) originally referred to Proconsul major are also now
included in Afropithecus (Leakey and Walker 1997).
Afropithecus fossils have been recovered from 21 separate sites
throughout the Kalodirr Member of the Lothidok Formation.
Originally dated to between 16 and 18 Ma on the basis of faunal
associations, two tuffs bracketing the Kalodirr Member have
now provided K/Ar ages for the Kalodirr Member. The Kalodirr
tuff, found immediately below the Kalodirr Member has been
dated to ca. 17.5 Ma and the capping Naserte tuff has been dated
to ca. 16.8 Ma (Brochetto et al. 1992). Additionally, the beds
from Morourot were assigned an age of ca. 17.5 Ma on the basis
of their position between the same radiometric ages (Brochetto
et al. 1992).
McDougall and Walthers (1985) reported an age of between
17.5 and 17.2 Ma for the hominoid-bearing beds of Buluk. These
beds span ten meters of a 120-meter section bracketed by K/Ar
dates of 18 and 17.2 Ma, with the fossil-bearing beds just below
the capping basalt. The fossil beds at Locherangan have been
dated to between 17.5 and 17 Ma (Anyonge 1991).
The early Miocene hominoid species Heliopithecus leakeyi, represented by a single maxillary fragment and isolated teeth, is
grouped by most workers with Afropithecus and Turkanapithecus
in the tribe ‘‘Afropithicini’’ (Andrews 1992; Andrews et al.
1996). The similarity of Heliopithecus to Afropithecus argues for
its inclusion of Heliopithecus within the Afropithecus OTU for this
study. Fossil remains of Heliopithecus have been recovered from
Ad Dabtiyah, Saudi Arabia (Andrews and Martin 1987b). The
age of Ad Dabtiyah is often cited as approximately 17 Ma (see
Andrews and Martin 1987b). Analysis of the vertebrate fauna at
the site (Gentry 1987a,b) more conservatively places Ad Dabtiyah between Rusinga (just younger than 18 Ma; see Proconsul)
and Maboko (younger than 16 Ma; see Equatorius).
DATA CONGRUENCE IN HOMINOID PHYLOGENY
Turkanapithecus
The genus Turkanapithecus is known only from the Kalodirr
locality in Kenya (Leakey and Leakey 1986b) and contains the
species Turkanapithecus kalakolensis. Recovered Turkanapithecus
material consists of 20 specimens including a nearly complete
cranium and mandible and several well-persevered postcranial
elements. Dating the Turkanapithecus material follows the dating
for the Kalodirr locality, but it was noted by Brochetto et al.
(1992) that the single Turkanapithecus site ‘‘directly underlies’’
the Naserte tuff (see Afropithecus above).
Equatorius
The genus Equatorius was only recently described by Ward et
al. (1999). By their definition all Kenyapithecus material not assigned to K. wickeri (known only from Fort Ternan, Kenya) is referred to the species Equatorius africanus. Equatorius is known
from several contemporary sites in the Maboko Formation, from
Kipsaramon in the Tugen Hills, and from Nachola in the Samburu Hills (Ward et al. 1999). The Maboko Formation sites (Kaloma, Ombo, Nyakach, Maboko, Majima) are estimated through
stratigraphic correlations to be between 15 and 16 Ma, on the
basis of their relative positions to radiometrically dated phonolitic units (Andrews 1981; Pickford 1985). The hominoidbearing beds of the Muryur Formation of the Tugen Hills are
slightly older than 15 Ma (Ward et al. 1999); they lie under the
Tiim Phonolite series dated (K/Ar) to between 15 and 13 Ma
(Hill et al. 1985). Recent dating (40Ar/39Ar) of the Muryur bed
has produced an age for the BPRP#122 locality of between 15.58
Ma and 15.36 Ma (Behrensmeyer et al. 2002). Specimens originally attributed to Kenyapithecus africanus from the Aka Aiteputh
and Nachola Formations in the Samburu Hills (Nakatsukasa et
al. 1998) were transferred to Equatorius by Ward et al. (1999).
These beds have been estimated to ca. 14.5 Ma (Nakatsukasa et
al. 1998). However, Ishida et al. (1999) have placed these fossils
in a separate taxon, Nacholopithecus kerioi, and the morphological
distinctness of this taxon from Equatorius was recognized by
Kelley et al. (2002). Awaiting a detailed analysis of this new taxon, we did not include Nacholopithecus in this study.
Kenyapithecus
With the separation of Equatorius africanus, Kenyapithecus is
here restricted to K. wickeri, known only from the ‘‘B’’ fossil
beds at Fort Ternan (Pickford 1985). The Baraget phonolitic lava
underlying the fossil-bearing unit has been dated by using K/
Ar to ca. 15 Ma (Shipman et al. 1981). K/Ar results on micas in
Fort Ternan paleosols give an age of 14 Ma with a hydrothermal
overprint in the paleosols of 13.5 Ma, implying that these units
had been deposited, pedogenically altered, and buried prior to
this date (Shipman et al. 1981). The Fort Ternan beds correlate
stratigraphically higher than Equatorius fossil localities (Ward et
al. 1999).
Griphopithecus
Griphopithecus alpani is known from the type locality at Çandır
and from Paşalar (both in Turkey) (Andrews et al. 1996). Currently a single species is recognized at Paşalar, although different tooth morphologies suggest two species are probably present (Alpagut et al. 1990). Fossils from the European localities
of Engelswies (cf. Griphopithecus) and Klein Haddersdorf, Austria, and Neudorf Sandberg (Griphopithecus cf. G. darwini) are included known (Andrews et al. 1996; Kordos 2000). It is not clear
if Griphopithecus material from these localities represents the
same taxon as the unnamed Paşalar species, and there is some
debate as to the assignment of the Engelswies material to this
genus (see below).
It appears that the hominoid-bearing unit at Paşalar is the result of a single flood event, possibly preserving an ecological
649
community (Andrews and Alpagut 1990; Bestland 1990). Paşalar is well studied, and faunal analyses have conventionally
placed the hominoid-bearing unit in the lower subzone of MN
6 on the basis of the appearance of rhinocerotids and proboscideans and analysis of cricetine rodents (Bernor and Tobien
1990). However, Peláez-Campomanes and Daams (2002) placed
Paşalar in later MN6 on the basis of a more complete analysis
of the rodent assemblage. The type locality for Griphopithecus, at
Çandır, had also been faunally correlated to MN 6 (Steininger
et al. 1996). Comparison of the faunas at these localities suggests
that Paşalar is slightly older than Çandır (Bernor and Tobien
1990). However, several biostratigraphic and magnetostratigraphic correlations place both the Çandır and Paşalar localities
within MN 5, ca. 16.5 Ma (Steininger 1999; Heizmann and Begun 2001). The European hominoid locality, Klein Haddersdorf,
also correlates to MN 6, and is believed to be younger than Paşalar (Begun 1992c; Heizmann and Begun 2001).
Magnetostratigraphic, lithostratigraphic, and biostratigraphic correlations to before the Langhian transgression, between
16.5 and 17 Ma indicate that Engelswies is the oldest evidence
for hominoids outside of Africa (Heizmann and Begun 2001).
However, Heizmann and Begun (2001) noted that hominoid material from Engelswies is tentatively assigned to Griphopithecus,
and possibly represents a distinct genus. With Çandır and Paşalar recalibrated to MN 5, the coding of the stratigraphic character for Griphopithecus is identical with or without the inclusion
of the Engelswies material in the OTU. Therefore, the FAE of
Griphopithecus is older than the western Kenyan Equatorius localities, and is penecontemporaneous with the LAE of Afropithecus.
Sivapithecus
The hominoid genus Sivapithecus is known from more than
100 fossil localities in the Siwalik molasse of India, Nepal, and
Pakistan. Although the dentition and mandible are well known
and the face is represented by a fairly complete specimen (GSP
15000 [Raza et al. 1983]), the postcranial anatomy of Sivapithecus
is rather poorly known. Several partial long bones and assorted
bones of the hand and foot are known; however, no remains of
the trunk have been recovered (Ward 1997b). The stratigraphy
of Sivapithecus is particularly well resolved. The commencement
of movement along the Main Central and Main Boundary
Thrust systems of the Himalaya Mountains is responsible for a
continuous flux of sediment into the Siwalik foreland for the last
18 million years, producing strata locally in excess of 4 km thick
with few hiatuses (N. M. Johnson et al. 1982; Opdyke et al. 1982;
Sorkhabi and Macfarlene 1999). Because much of the Siwalik sequence has been calibrated to the GPTS (G. D. Johnson et al.
1982), Sivapithecus can be accurately placed into a worldwide
chronology. Kappelman et al. (1991) placed the FAE of Sivapithecus at the Y750 locality near Chinji, Pakistan, which has been
paleomagnetically dated to Chron 5Ar.1 of the GPTS (12.68–
12.71 Ma [Cande and Kent 1995]). Speculation over the presence
of Sivapithecus older than 13 Ma (the GSP 15000 face [Raza et al.
1983]) was discounted with a stratigraphic reassessment placing this locality stratigraphically higher than Y750 (Kappelman
et al. 1991). The LAE of Sivapithecus is likely recorded at the GSID-185 locality near Haritalyangar, India (Patnaik and Cameron
1997). The paleomagnetic results of Johnson et al. (1983) estimate this site to be ca. 7.4 Ma in age, although some disagreement exists over the correlations at Haritalyangar (Kelley and
Pilbeam 1986).
Ankarapithecus
Hominoid fossils from the locality of Yassıören, Turkey, were
originally known from two gnathic fragments with teeth (Ozansoy 1965) and a palate with the right zygomatic arch (Andrews
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JOHN A. FINARELLI AND WILLIAM C. CLYDE
and Tekkaya 1980). Both were found at Locality 8A in the Sinap
Formation and assigned to the taxon Sivapithecus meteai (Andrews and Tekkaya 1980). Restoration and reevaluation of the
palate led to the separation of this material from Sivapithecus
(Begun and Güleç 1995). In 1996, a face and unassociated mandible were recovered from Locality 12 at Yassıören (Alpagut et
al. 1996). Although separated by over a kilometer, localities 8A
and 12 correlate stratigraphically to within a few meters of one
another and are considered contemporaneous. Paleomagnetic
surveys of the Sinap Formation at Yassıören place the hominoidbearing beds in Chron C5n.1n of the GPTS (Alpagut et al. 1996).
Recent paleomagnetic surveys of the Vallès Penedès, Spain, correlate Ankarapithecus as contemporary with the MN 9 Zone for
western Europe (Krijgsman et al. 1996; Agustı́ et al. 1998, 2001),
indicating that Ankarapithecus was a contemporary of both Sivapithecus and Dryopithecus (see below).
Dryopithecus
Currently four species are recognized in the genus Dryopithecus. Dryopithecus fontani is known from St. Gaudens, France (Begun 1994), and from St. Stephan, Austria (Andrews et al. 1996).
In eastern and northern Spain, the taxa D. laietanus and D. crusafonti have been recovered from seven localities in the Vallès
Penedès and from Seu d’Urgell (‘‘El Firal’’). Dryopithecus brancoi
is known from several sites in Eastern Europe, including Salmendigen, Germany (type) and Rudabanya, Hungary (Begun
and Kordos 1993).
The sites in the Vallès Penedès have been correlated to Zones
MN 8 (San Quirze, Can Vila) through MN 10 (La Tarumba) (Begun et al. 1990). Recent paleomagnetic surveys in the Vallès Penedès have restricted the correlations of these sections to MN 9
(Chron C5r.1r: 11.1 Ma) to middle MN 10 (ca. 9.2 Ma), with the
site of Can Llobateres straddling the MN9/10 transition at ca.
9.7 Ma (Agustı́ et al. 2001). This is in agreement with the MN 9
faunal age given to the Seu d’Urgell, which has been correlated
to Can Llobateres (Begun 1992a). Salmendigen is correlated to
the earliest part of MN 10 (Begun 1994), roughly equivalent to
La Tarumba, whereas Rudabanya, has been correlated to MN 9
on the basis of the mollusk and vertebrate faunas (Begun 1992c).
St. Stephan is correlated to MN 8 (Andrews et al. 1996). The material from St. Gaudens presents the most troublesome data for
Dryopithecus. The locality is not well sampled, and the age is correspondingly not well known. Begun (1992c) places the site at
latest MN 7 or, more probably, early MN 8.
Ouranopithecus
Ouranopithecus ( 5 Graecopithecus [see de Bonis and Koufos
1993 for summary of the debate]) is known from three fossil localities in Greece. Two of these localities, Ravin de la Pluie and
Xirochori, lie in the Nea Messimbria Formation. Multiple maxillary and mandibular remains have been recovered from Ravin
de la Pluie, and Xirochori has yielded a distorted face (de Bonis
and Koufos 1993). The third locality lies within the Nikiti Formation and has produced a mandible and a female maxilla
(Koufos 1995).
Faunal analysis of Ravin de la Pluie and Xirochori indicate
that this site is in MN Zone 10 (latest Vallesian Land Mammal
Age [de Bonis and Koufos 1993]). A paleomagnetic survey confirms this, correlating the overlying formation to Chron 8 and
MN Zone 11 (Kondopulou et al. 1992). Faunal analysis of the
locality at Nikiti gives an age of MN 10 to MN 11 (Koufos 1995).
It should be added that an additional specimen from Pyrgos,
Greece, unearthed during the construction of a swimming pool,
purports to extend the range of Ouranopithecus to beyond MN
Zone 11 on the basis of the presence of Hipparion. The association of the Hipparion fossils, however, has been called into question and the locality was destroyed during local construction
(de Bonis and Koufos 1993). Therefore, Ouranopithecus is limited
in this study to the three sites for which verifiable stratigraphic
data are available, and it is treated here as ranging from latest
MN 10 through MN 11.
Lufengpithecus
Lufengpithecus is a late Miocene hominoid from the Shihuiba
colliery site 9 km north of Lufeng, China. Although Lufengpithecus lufengensis is the only species formally named for this genus, some debate exists as to whether larger and smaller specimens recovered from this locality represent males and females
in a highly dimorphic ape (such as Pongo), or if they represent
two congeneric species (see Kelly and Plavcan 1998). Faunal
analysis of rhyzomyid rodents at the site are originally published in Chinese (Wu et al. 1986) and are generally quoted as
ca. 8 Ma in the English literature (Kelly and Plavcan 1998;
Schwartz 1997), although they may indicate an age as young as
7 Ma (see Badgely et al. 1988). Pilbeam et al. (1996) however,
correlated Lufeng biostratigraphically to the Dhok Pathan Formation in the Siwaliks, roughly 9 Ma. An age of approximately
8–9 Ma correlates Lufengpithecus to near the European MN Zone
11/12 transition (Harrison et al. 2002; Steininger et al. 1996),
and thus with the lower (V1) portion of the Oreopithecus range
(see below). Additional material from the Xiaohe Formation in
the Yaunmou Basin correlates biostratigraphically to just older
than the Shihuiba locality near Lufeng (Harrison et al. 2002).
Reports of hominoid material have been reported from the Yangyi locality, although these fossils have yet to be completely described. Dates for this locality have been reported at roughly 4
Ma on the basis of proboscideans, although these dates are, as
yet, not reliably established (Harrison et al. 2002). As such, the
discussion of Lufengpithecus here will incorporate the Shihuib
and Yaunmou localities for which there is formally described
material and the stratigraphy is reasonably well constrained.
Oreopithecus
The hominoid genus Oreopithecus is represented by a single
species, O. bambolii. Oreopithecus has been recovered from five
localities in the Maremma Valley in southwestern Tuscany (Azzaroli et al. 1986), a lignite bed in Serrazzano, Italy, and one locality in Sardinia (Harrison and Rook 1997). Early work divided
the Maremma localities into three faunal horizons: V1, V2 and
V3 (Delson and Szalay 1985). Oreopithecus material has been recovered from the lower V1 and V2 faunas. Hominoid material
also has been recovered from the V1/V2 intermediate ‘‘Cardium Horizon’’ at Baccinello (Harrison and Rook 1997). Baccinello is the only locality where Oreopithecus has been documented spanning both horizons. The Serrazzano and Sardinia
localities correlate to the V2 horizon of Baccinello (Rook et al.
1996).
A recent 40Ar/39Ar date of a felsic tuff in the V2 horizon at
Baccinello gives an age of 7.55 6 0.03 Ma (Rook et al. 2000). The
V1 horizon is known from 150 m below this date, although sedimentation rates are thought to be high at Baccinello, so the absolute time represented by the section is not thought to be extensive. Faunal analysis, specifically the presence of Huerzelermys vireti ( 5 ‘‘Valerimys aff. vireti’’ of Huerzeler and Engesser
1976), correlates the V1 and V2 horizons to the early Turolian.
The Cardium Horizon at Baccinello has produced an MN 12 and
MN 13 intermediate fauna. No paleomagnetic information has
been published for the Oreopithecus localities. Revised dating of
the Neogene Mammal Zones (Steininger et al. 1996; Agustı́ et
al. 2001) agrees with the radiometric age obtained for Baccinello
(Rook et al. 2000). Therefore, a correlation of the V1 horizon
with latest MN 12 and the V2 horizon with earliest MN 13 is
used here.
DATA CONGRUENCE IN HOMINOID PHYLOGENY
Australopithecus
Australopithecus is used here as an OTU to represent hominins
(5 human and all direct ancestors [Tattersall et al. 1988]). The
oldest known fossils of this genus are from Kanapoi, Kenya, attributed to the species A. anamensis (Leakey et al. 1995a). Ar/Ar
ages on bracketing tephra place Kanapoi between 4.17 and 4.12
Ma (Leakey et al. 1999). From this point, the hominin lineage
has a fairly continuous fossil record through the Recent. An older fossil hominin from Aramis, Ethiopia, originally assigned to
‘‘Australopithecus’’ ramidus (White et al. 1994) but later renamed
Ardipithecus ramidus (White et al. 1995) has not yet been fully
described in the literature, although potential evidence of bipedality has been noted, indicating a link with later hominins.
The Aramis VP Locality 6 directly overlies the Gàala Vitiric Tuff
Complex (White et al. 1994), which has been dated by 40Ar/39Ar
to 4.39 Ma (WoldeGabriel et al. 1994). The FAE of the OTU Australopithecus can be defined using either A. ramidus or A. anamensis without creating overlap with Oreopithecus.
Recent discoveries of potential hominins Orrorin tugenensis
(Senut et al. 2001) and Sahelanthropus tchadensis (Brunet et al.
2002) will likely have profound effects on our understanding of
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early human evolution. Orrorin is known from the Lukeino Formation of the Tugen Hills and has been dated to ca. 6 Ma (Senut
et al. 2001). If Orrorin is the direct ancestor to the modern human lineage as proposed by Senut et al. (2001), this date would
not affect the stratigraphic character. However, if Australopithecus represents an evolutionary ‘‘side branch,’’ then the use of the
OTU Australopithecus as a proxy for the human lineage (as in this
study and Begun et al. 1997) would have to be reexamined carefully. However, Aiello and Collard (2001) urged caution in accepting the hypothesis that Orrorin is the direct ancestor to
modern Homo, and the use of Australopithecus as a representative
for the hominin lineage is retained here. Sahelanthropus was recently described form the Toros-Menalla locality TM 266, in the
western Djurab Desert of northern Chad. The fossil-bearing anthracotheriid unit has been tentatively dated to between 6 and
7 Ma on the basis of biochronological correlations with Lukeino
and Lothagam (Brunet et al. 2002). As with Orrorin, this age
would have no effect on the coding of the stratigraphic character
if Sahelanthropus represents an early member of the hominin
clade. However, a cladistic analysis of the Sahelanthropus material is, as yet, not possible.