Journal of Human Evolution 57 (2009) 697–709
Contents lists available at ScienceDirect
Journal of Human Evolution
journal homepage: www.elsevier.com/locate/jhevol
New primate carpal bones from Rudabánya (late Miocene, Hungary):
taxonomic and functional implications
Tracy L. Kivell*, David R. Begun
Department Anthropology, University of Toronto, 19 Russell Street, Toronto, ON M5S 2S2, Canada
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 4 July 2008
Accepted 19 May 2009
We describe a scaphoid and two capitates from the late Miocene site of Rudabánya, Hungary using
qualitative and quantitative comparisons to a large sample of hominoid, cercopithecoid, and platyrrhine
primates. The scaphoid (RUD 202) is not fused to the os centrale and in this way is like most primates
other than African apes and humans (hominines). Qualitatively, its morphology is most similar to Pongo,
and univariate analyses generally confirm an ape-like morphology with an increased range of mobility.
One capitate (RUD 167) is compatible in size to the scaphoid, and its morphology suggests a combination
of monkey-like generalized arboreality and ape-like enhanced mobility. RUD 203 is a smaller, fragmentary capitate, about half the size of RUD 167, and preserves only the distal portion of the body with
the third metacarpal articular surface. Its morphology is virtually identical to that of RUD 167, and an
exact randomization test revealed that it is statistically likely to find two carpal bones of such disparate
sizes within one taxon. However, due to morphological similarities with other Miocene hominoids as
well as implications for size variation within one taxon and sex, we consider the taxonomic affiliation of
RUD 203 to be unresolved. We attribute the scaphoid and RUD 167 capitate to the hominine Rudapithecus
hungaricus (formerly Dryopithecus brancoi; see Begun et al., 2008) based on overall morphological
similarity to extant apes, particularly Pongo, and not to the pliopithecoid Anapithecus hernyaki, the only
other primate known from Rudabánya. The similarities in carpal morphology to suspensory taxa are
consistent with previous interpretations of Rudapithecus positional behavior. The scaphoid and the RUD
167 capitate are consistent in size with a partial skeleton including associated postcranial and craniodental specimens from the same level at the locality and may be from the same individual. These are the
first carpal bones described from Rudabánya and from this taxon, and they add to our understanding of
the evolution of arboreal locomotion in late Miocene apes.
Ó 2009 Elsevier Ltd. All rights reserved.
Keywords:
Capitate
Scaphoid
Wrist
Suspensory locomotion
Rudapithecus
Dryopithecus
Anapithecus
Introduction
This report describes three fossil primate carpal bones, a scaphoid
and two capitates, recovered from the late Miocene (10 Ma) site of
Rudabánya in northern central Hungary (Kordos and Begun, 2002).
These are the first primate carpal bones to be described from
Rudabánya (Begun et al., 2003; Kivell and Begun, 2006), and thus
they hold unique taxonomic and functional information about
primate evolution during this time period. There are currently two
primate taxa recognized at Rudabánya: Anapithecus hernyaki and
Rudapithecus hungaricus (Begun et al., 2008; Kordos and Begun,
2001a). Anapithecus is a roughly 10 to 15 kg pliopithecoid, a primitive
catarrhine (Begun, 2002). The few postcranial remains attributed to
Anapithecus are broadly similar morphologically to Epipliopithecus
from Slovakia and indicate a primate that is generally well-adapted
to an arboreal environment (Begun, 1988, 1993).
* Corresponding author.
E-mail address: [email protected] (T.L. Kivell).
0047-2484/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2009.05.011
Rudapithecus is a fossil great ape with synapomorphies of
extant African apes, which we include in the Homininae (Begun,
1994, 2007; Begun and Kordos, 1997; Kordos and Begun, 2001b).
Previously the fossil hominine (defined as the clade including
African apes, humans, and their ancestors) from Rudabánya was
assigned to the taxon Dryopithecus brancoi (Begun and Kordos,
1993). Recent analysis of the large number of relatively complete
craniodental and postcranial specimens have led us to conclude
that the dryopithecini are more diverse at the genus level (Begun
et al., 2008; Begun, in press), and thus we elevate the genus
Rudapithecus (Kretzoi, 1969) from junior subjective synonym
status with Dryopithecus.1 Functional interpretations of
1
The rationale for this taxonomic decision is outlined in Begun (in press). It is
consistent with the recent recognition of Hispanopithecus for the sample from Can
Llobateres which has long been described as Dryopithecus laietanus (Begun, 2002;
Almecija et al., 2007).
698
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
postcranial remains indicate that the positional behavior of
Rudapithecus included suspension, climbing, and some quadrupedalism (Morbeck, 1983; Begun, 1988, 1992, 1993, 1994; Kivell
and Begun, 2006). Body mass estimates for Rudapithecus range
between roughly 20–40 kg (Morbeck, 1983; Begun, 1994),
suggesting a large degree of sexual dimorphism similar to that of
extant Pongo or Gorilla.
Among the primate postcranial remains uncovered at Rudabánya are three carpal bones. The RUD 202 scaphoid was recovered
in 1993 from the same layer of the R. II site at Rudabánya as other
specimens attributed to a skeleton, recovered between 1998 and
2006 (Kordos and Begun, 1997, 2001a,b, 2002). The scaphoid was
recovered with the distal one third of a metacarpal. Both specimens have the same preservation and are of comparable size,
suggesting they are from the same individual (Zylstra et al., 2007).
Both capitates were found in the museum collections, from excavations conducted in the 1970s. The nature of the preservation and
the matrix indicates that the specimens also come from the same
level of R. II as the scaphoid, metacarpal, and partial skeleton (i.e.,
the ‘‘grey marl’’ [Kordos and Begun, 2001a]). Museum records
indicate that this level was sampled in the year in which these
specimens were found. The larger of the two capitates may be
associated with the same individual as the scaphoid, while the
other specimen is too small to be from the same individual.
Continued excavations at Rudabánya will be directed at assessing
the taphonomy of these specimens and additional postcranial
material and at the likelihood that all but the small capitate belong
to the same individual.
RUD 202 is a right scaphoid. The body is largely complete,
missing only the dorsal edge of the radial facet, and the scaphoid
tubercle is broken at the ‘neck.’ RUD 167 is a nearly complete right
capitate of appropriate size for the scaphoid. Slight abrasion occurs
on the lateral, dorsal, and palmar surfaces of the proximal facet, or
capitate ‘head,’ and there is slight damage to the trapezoid/second
metacarpal articular area and dorsal edge of the hamate facet. RUD
203 is also a right capitate but is smaller overall than that of RUD
167. Only the distal portion of the capitate with the third metacarpal articular surface is preserved.
The aims of this analysis are to (1) describe the functional
morphology of the carpal bones and (2) determine the taxonomic affiliation of each to either Rudapithecus or Anapithecus
(or neither) as recognized at Rudabánya. Other postcranial fossils
attributed to the dryopithecini, specifically Rudapithecus, show
distinct morphological features indicative of fore- and hind limb
suspension and suspensory locomotion (Morbeck, 1983; Rose,
1983; Begun, 1988, 1992). Thus, we hypothesize that if a carpal
bone is attributed to Rudapithecus it should show hominoid-like
morphology with features reflective of increased mobility. The
paucity of postcranial remains attributed to Anapithecus does not
permit a detailed reconstruction of the locomotor behavior of
this primitive catarrhine nor the formulation of clear predictions
of the most likely carpal morphology. Therefore, we have chosen
a comparative sample that will cover all likely patterns of
positional behavior. Variation in size may also help to further
discriminate between the larger Rudapithecus and smaller Anapithecus, although there may be some overlap between these
two taxa since the degree of sexual dimorphism in each is
unknown. If the morphology of these fossils does not fit confidently in either of these taxonomic groups (or even if they do), it
is, of course, possible that these carpal bones come from
a different primate not yet recognized at Rudabánya. We
compare these fossil carpal bones to a sample of fossil and
extant apes and monkeys of a variety of locomotor behaviors in
order to interpret the functional morphology and to test these
predictions.
Materials and methods
We analyzed the morphology of all three carpals both qualitatively and quantitatively. We made quantitative comparisons to
a large sample (n ¼ 227) of adult extant hominoids, cercopithecoids,
and platyrrhines (Table 1a) and to Miocene hominoids Proconsul,
Afropithecus, Oreopithecus, and Sivapithecus (Table 1b). Species were
grouped primarily by locomotor behavior as well as broad taxonomic categories to best quantify the variation in morphology and
to provide the most meaningful comparisons. The species included
in each group are listed in Table 1a. This comparative sample was
chosen because it encompasses a broad range of haplorhine
primates and locomotor behaviors that are most informative for
interpreting the functional morphology of the primate taxa already
found at Rudabánya. New World monkeys may be a particularly
informative comparison because (1) morphological similarities of
the forelimb have been previously documented between Miocene
hominoids and platyrrhines (e.g., Rose, 1983) and, (2) cercopithecoids likely passed through a terrestrial phase during their evolution
and, as a consequence, almost certainly do not represent the
primitive crown catarrhine condition (Andrews, 1982).
Six morphometric variables on the scaphoid and eight variables
on the capitate quantified the overall size of each carpal and size of
the articular facets (Fig. 1, Table 2). All measurements were taken on
the original fossil specimens of the Rudabánya carpal bones, but
comparisons to other Miocene hominoids were taken from casts.
Since body mass is not available for fossils or most extant specimens
in osteological collections, each value was divided by a geometric
mean to adjust for size (Mosimann and James, 1979; Jungers et al.,
1995). Due to the fragmentary preservation of capitate RUD 203, the
geometric mean was derived from only two variables (height and
breadth of the capitate body). Since the geometric mean is a volume
that requires at least three variables (Coleman, 2007), we tested that
a geometric mean derived from just two variables was significantly
correlated with the geometric mean derived from all eight capitate
variables. Spearman rank correlation was used and a significant
correlation was determined as rs > 0.80 (Quinn and Keough, 2002).
For the extant sample, differences in carpal variable means were
examined with analysis of variance (ANOVA) followed by a TukeyKramer post-hoc test (Quinn and Keough, 2002). All statistical
analyses were run with sexes pooled. Differences in carpal variables
among extant groups and the fossil sample were evaluated graphically with box-and-whisker plots.
Given the variation in size between the two capitates, we tested
the null hypothesis that both specimens were from a single species
using two methods. First, we measured the average degree of sexual
Table 1a
Comparative sample of adult (a) extant haplorhines and (b) Miocene hominoid
carpal sample.
Taxon
Locomotion
Total n
_
\
Pan paniscus
Pan troglodytes
Gorilla
Pongo
Hylobates
Symphalangus
Papio
Macaca mulatta
arboreal knuckle-walking
arboreal knuckle-walking
terrestrial knuckle-walking
suspensory
brachiating
brachiating
cercopithecoid terrestrial quadruped
cercopithecoid semi- terrestrial
quadruped
cercopithecoid semi-terrestrial
quadruped
cercopithecoid arboreal quadruped
21
32
45
32
38
5
10
12
10
16
26
13
17
3
5
6
11
16
19
19
21
2
5
6
7
4
3
8
4
4
New World monkey suspensory
New World monkey arboreal
quadruped
4
13
3
3
1
10
Chlorocebus
Macaca
fascicularis
Ateles
Alouatta
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Table 1b
Taxon
Carpal
Specimen
Proconsul heseloni
scaphoid
capitate
scaphoid
scaphoid
capitate
capitate
capitate
capitate
scaphoid
capitate
capitate
capitate
scaphoid
KNM-RU 2036
KNM-RU 2036
C14
C11/50
C26
C25
C28
KNM-RU 1907
KNM-SO 999
KNM-CA 409
KNM-WK 18365
GSP Y500 17119
Basel 36
Proconsul africanus
Afropithecus turkanensis
Sivapithecus indicus
Oreopithecus bambolii
dimorphism in extant taxonomic groups. We calculated indices of
sexual dimorphism (ISD) for each extant taxon and between the pair
of fossil capitates to determine the likelihood that the size variation
between the fossils was reasonable for opposite sexes of the same
species (Lockwood et al., 1996). The ISD is the ratio of the male to
female mean values of the geometric mean (Lockwood et al., 1996).
Second, we used exact randomization to test the likelihood of
finding two individuals from the same extant population with the
same or greater degree of size variation as found between the two
fossil capitates (Richmond and Jungers, 1995). We calculated the
difference (Di) between geometric means for every possible pairwise comparison within each species, including opposite- and
same-sex pairs (Richmond and Jungers, 1995; Quinn and Keough,
2002). The proportion of Dis that were greater or equal to the
difference between the original mean (D0 or the ISD) is the p-value
(alpha ¼ 0.05) and determines the likelihood of sampling two
individuals as disparate in size as that observed between the fossil
pair (Quinn and Keough, 2002).
Results
All three carpal bones are adult in morphology. Despite some
damage, each carpal preserves portions of the body that are last to
fully ossify during ontogeny, and facets are well defined, as is only
found in adult carpal morphology (Kivell, 2007). All three specimens are broadly within the size range of Papio and are smaller
than extant great apes. Raw mean values, standard deviations, and
the range for each morphometric variable are given in Table 3 for
the scaphoid and in Table 4 for the capitate.
Scaphoid RUD 202
The scaphoid is similar in size to Old World monkeys that are
within the body mass range of about 16 to 22 kg (e.g., Papio, Nasalis,
Presbytis). Qualitatively, RUD 202 is most similar in morphology to
Pongo and distinctly different from that of African apes. The overall
shape of the body is distolaterally elongated and is not fused to the
os centrale, as in most non-hominine primates (Fig. 2). The tubercle
and trapezium facet are not preserved. The scaphoid is waisted and
broad at the neck and indicates an elongated, more distally-projecting tubercle; a composite of features that is typical of Asian
apes. The tubercle, however, appears slightly more palmarlyoriented than that of the highly-suspensory Oreopithecus. This
morphology is different from the narrow tubercle base and often
more palmarly-oriented tubercle seen in African apes. The lunate
facet is Asian ape-like in its large size and distal elongation to the
base of the tubercle and is most similar to Pongo and hylobatids in
this respect (though it is not as dorsopalmarly tall as in Pongo or
Oreopithecus). Again, the lunate facet morphology is distinctly
699
different from the small, proximally-confined lunate facet of
African apes. The os centrale facet is small and circular, like that of
monkeys, and lacks the distal extension of this facet that is typical
of Asian apes. The os centrale articular surface is more dorsally
angled relative to the lunate surface, most similar to Asian apes, and
unlike the almost coplanar surfaces found in Oreopithecus. Damage
to the proximal edge of the scaphoid makes it difficult to interpret
the overall shape of the radial facet. However, the preserved
portion is more proximally rounded than that of Proconsul and is
more monkey or Asian ape-like in the overall shape of the facet. The
radial facet is extensive, particularly dorsally, and, again, appears
most similar to Pongo.
Univariate analysis of the size-adjusted scaphoid variables
generally supports the qualitative assessment of the morphology.
The RUD 202 scaphoid is hominoid-like, unique from that of other
Miocene taxa, and shares clear similarities to great apes and more
arboreal apes, in particular (Fig. 3). ANOVA reveals significant
differences among extant taxonomic-locomotor groups for all
variables. The RUD 202 scaphoid is most similar to great apes and
suspensory platyrrhines in relative height of the body and especially similar to suspensory hominids (Pongo) in its breadth
(Fig. 3a, b). The shape of the radial and lunate facets show the
clearest functional signal and are thus most informative for
interpreting fossil morphology. Compared to other Miocene taxa,
RUD 202 has the most extensive radial facet and an extremely
long lunate facet (Fig. 3c, d, f). The relatively tall radial facet of
RUD 202 is most similar to cercopithecoids and knuckle-walking
apes but is within the range of all taxa except platyrrhines
(Fig. 3c). The relatively long length of the radial facet is most
similar to arboreal knuckle-walkers and unlike suspensory or
brachiating taxa (Fig. 3d). The RUD 202 lunate facet is more
distally elongated than any other fossil or extant taxon and is
most similar to suspensory and brachiating taxa. It is important to
note that semi-terrestrial and arboreal cercopithecoids also have
relatively long lunate facets. However, their morphology is
different from that of all other non-cercopithecoid taxa (including
RUD 202), such that the lunate facet extends onto the medially
oriented tubercle.
Capitate RUD 167
RUD 167 is consistent in size with the scaphoid RUD 202, suggesting the same range of body size. The overall shape of the capitate
is that of a generalized arboreal primate, lacking the stabilizing
adaptations of African apes and terrestrial monkeys or the adaptations to increased midcarpal mobility seen in Asian apes (Begun,
2004; Richmond, 2006; Fig. 4). In dorsal view, the capitate body is
not strongly ‘‘waisted’’ as in African apes and some monkeys,
although damage to the head may exaggerate this impression. RUD
167 is more similar to arboreal monkeys, Proconsul, and to
morphology reported for Dryopithecus (¼Pierolapithecus; Moyà-Solà
et al., 2004), lacking mediolateral expansion of the head and distal
portion that is typical of great apes and terrestrial monkeys. The
articulation for the os centrale along the lateral side is uniquely
expansive, in that it is extended distally along both the dorsal and
palmar portions. The palmar extension of this facet is most similar to
hylobatids and unlike the palmarly-truncated facet of great apes and
monkeys. The distal portion of the lateral side preserves a distinct
circular articulation palmarly and a smaller facet dorsally for the
second metacarpal (MC2), both of which are oriented laterally. The
separation between these facets is unlike the continuous articulation
typical of most monkeys and some Proconsul specimens and is most
similar to great apes and Papio. RUD 167 facet shapes are unique
compared to our extant sample, with the closest comparison being
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T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Figure 1. Morphometric variables of the scaphoid and capitate. (a) Scaphoid shown in distomedial (above), proximolateral (below), and palmar (right) views. (b) Capitate shown in
dorsal (left) and medial (right) views. See Table 2 for abbreviations.
with Papio. An articulation for the trapezoid, just proximal to the
dorsal MC2 facet, is not clear due to abrasion but was likely present,
as in most primates (but unlike Gorilla; Lewis, 1973).
The distal, third metacarpal (MC3) articular surface is most
similar to Proconsul and extant monkeys in its general outline, with
a ‘squared’ palmar half that is unlike the palmar, lateral extension of
Asian apes. However, the facet morphology is generally more
similar to hominoids in that the metacarpal surface is slightly
keeled with a well-defined vertical ridge that divides the articular
surface into two distinct facets; a larger, medial portion that is
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Table 2
Description of morphometric variables in the scaphoid and capitate
Scaphoid
Description
HSB
BSB
LSRF
HSRF
LSLF
HSLF
max.
max.
max.
max.
max.
max.
Capitate
Description
LCB
HCB
BCB
HCHF
LCHF
BCPF
HCPF
BCN
max proximodistal length of capitate body
max dorsopalmar height of capitate body
max mediolateral breadth of capitate body (in distal view)
max dorsopalmar height of capitate hamate facet
max proximodistal length of capitate hamate facet
max mediolateral breadth of capitate proximal facet
max dorsopalmar height of capitate proximal facet
min mediolateral breadth of capitate neck
dorsopalmar height of scaphoid body
mediolateral breadth of scaphoid body (excl. tubercle)
dorsopalmar length of scaphoid radial facet
proximodistal height of scaphoid radial facet
dorsopalmar length of scaphoid lunate facet
proximodistal height of scaphoid lunate facet
slightly concave and oriented at a slight medial angle and a smaller,
lateral portion that is restricted to the dorsal half of the capitate
body and oriented slightly laterally. The MC3 facet morphology in
RUD 167 is similar to that of Sivapithecus and Proconsul and is unlike
the more irregular, keeled surfaces of extant African apes or the
single, concave articular surface typical of monkeys.
The hamate facet dominates the medial side of the RUD 167
capitate, extending the proximodistal length of the dorsal portion
of the capitate body, but is damaged at the midline. The hamate
facet morphology is unremarkable and broadly similar to Proconsul
or Pan in its shape. The articular surface is only slightly concave as
in Pongo, arboreal monkeys, and especially Proconsul and unlike the
stronger curvature found in terrestrial monkeys and African apes.
The palmar half of the capitate body has a small but distinct
medially-oriented facet of the fourth metacarpal that is more
monkey-like and is not found in most extant hominoids.
701
Univariate analysis of the size-adjusted capitate variables
supports the qualitative assessment of RUD 167 morphology
(Fig. 5). The overall shape of the RUD 167 capitate body is most
similar to suspensory and brachiating hominoids and arboreal
knuckle-walkers (Fig. 5a–c). While length of the hamate facet
appears more generalized (Fig. 5d), the height is relatively tall
compared to most other Miocene taxa and all extant taxa and is
most similar to suspensory and brachiating hominoids (Fig. 5e). The
shape of the proximal facet yielded the strongest functional signal
among extant taxa (Fig. 5f, g). RUD 167 has a relatively small
proximal facet compared to other Miocene taxa, overlapping with
only some P. heseloni specimens (KNM-RU 2036 in relative height
and KNM-RU 1907 and C25 in relative breadth). Compared to
extant taxa, RUD 167 is most similar to suspensory apes and
arboreal quadrupedal monkeys (both cercopithecoids and platyrrhines). The breadth of the capitate neck was similar to most other
Miocene taxa and, again, closest to suspensory apes among the
extant taxa (Fig. 5h).
Capitate RUD 203
The RUD 203 capitate is similar in size to Ateles or P. heseloni,
suggesting a body size of about 10 kg. Despite this smaller size
relative to RUD 167, both capitates share almost identical
morphology of the distal metacarpal facets (Fig. 4). RUD 203 also has
a well-defined ridge that demarcates a smaller lateral portion that is
oriented more laterally, from a larger medial articular area. Both facet
areas are relatively smooth and slightly concave and differ from the
more irregular surface typical of great apes. Facets for the MC2 are
preserved on the lateral side and are, again, very similar to RUD 167;
the palmar facet is well-demarcated and circular and is separated
from the dorsal facet as in extant hominoids. As in RUD 167, poor
preservation makes it difficult to interpret the presence and
morphology of a trapezoid facet.
Table 3
Scaphoid morphometric variables in fossil taxa and extant locomotor-taxonomic groupsa
Locomotor-taxonomic group
Other Fossils
Oreopithecus n ¼ 1
Proconsul africanus n ¼ 1
Proconsul heseloni n ¼ 5
RUD 202 n ¼ 1
Cercopithecoid
Arboreal quadruped n ¼ 8
Semi-terrestrial n ¼ 19
Terrestrial quadruped n ¼ 10
New World monkey
Arboreal quadruped n ¼ 12
Suspensory n ¼ 4
Hominoid
Brachiator n ¼ 43
Suspensory n ¼ 32
Arboreal knuckle-walker n ¼ 53
Terrestrial knuckle-walker n ¼ 45
a
HSB
BSB
HSRF
LSRF
HSLF
LSLF
13.80
10.33
10.23 (0.11)
10.11–10.32
8.45
5.73
6.36 (0.52)
5.95–6.95
13.63
9.62
9.97 (0.94)
8.89–10.55
14.10
10.35
9.80 (0.84)
9.04–10.71
9.57
7.37
5.85 (0.21)
5.63–6.05
6.72
6.16
7.25 (1.64)
5.36–8.37
11.82
8.04
11.26
11.66
7.08
13.11
6.14 (0.64)
5.22–7.32
7.87 (1.81)
5.39–10.21
12.24 (1.6)
9.57–14.32
6.25 (0.56)
5.38–6.92
7.84 (1.78)
5.12–10.19
11.76 (1.21)
10.41–13.5
7.08 (0.61)
5.96–7.71
8.83 (1.88)
5.46–12.19
12.75 (2.18)
9.93–16.58
6.10 (0.70)
5.06–7.25
7.65 (1.3)
5.59–10.13
12.18 (1.85)
9.47–14.71
4.06 (0.80)
2.94–5.06
4.37 (0.72)
3.27–6.15
7.06 (1.19)
5.43–9.36
6.14 (1.31)
4.08–8.03
7.53 (1.55)
4.44–10.80
9.83 (1.88)
6.96–13.29
11.87 (0.52)
11.05–12.67
10.29 (3.58)
7.4–15.21
4.28 (0.33)
3.80–4.86
6.03 (0.44)
5.71–6.68
6.94 (0.31)
6.55–7.41
8.44 (0.63)
7.89–9.18
7.05 (0.45)
6.17–7.70
9.06 (1.44)
7.44–10.84
5.27 (0.71)
4.19–6.76
6.06 (1.12)
4.72–7.04
5.68 (0.60)
4.85–7.04
7.84 (1.33)
6.41–9.01
8.40 (0.49)
7.26–9.42
17.24 (2.63)
13.09–23.92
18.36 (1.91)
15.15–23.90
24.32 (3.74)
18.87–38.96
6.36 (1.00)
4.56–9.74
13.11 (2.61)
8.37–19.27
11.89 (1.56)
9.31–18.3
15.41 (2.55)
11.62–21.96
8.03 (1.36)
5.45–10.36
16.37 (1.58)
13.72–19.23
18.06 (2.33)
14.07–23.49
23.11 (3.07)
18.25–30.86
8.76 (0.87)
6.20–10.90
16.02 (2.43)
12.40–20.64
17.46 (1.83)
13.08–22.46
21.49 (3.04)
15.17–30.26
5.98 (0.67)
4.66–7.55
9.44 (2.24)
6.26–15.25
10.78 (1.42)
8.44–13.78
15.50 (2.56)
11.29–21.45
7.80 (0.95)
5.61–10.46
15.20 (2.14)
12.23–22.07
8.73 (1.67)
5.62–13.53
12.58 (2.32)
7.08–17.46
See Table 2 for variable abbreviations. For each group, mean values (with standard deviation) are listed in the first line, and the range of values is listed in the second line.
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T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Table 4
Capitate morphometric variables in fossil taxa and extant locomotor-taxonomic groupsa
Locomotor-taxonomic group
LCB
HCB
BCB
HCHF
LCHF
BCPF
LCPF
BCN
Sivapithecus n ¼ 1
16.70
12.35
13.22 (1.83)
10.09–14.80
21.92
15.13
9.19
9.72 (1.45)
7.15–10.52
20.65
10.82
7.30
7.61 (1.2)
5.71–8.42
12.81
9.02
6.41
6.19 (0.45)
5.81–6.90
14.53
14.52
10.61
11.17 (2.02)
7.65–12.54
17.65
7.48
5.04
5.43 (0.99)
3.99–6.34
10.45
9.75
6.11
6.50 (1.09)
4.64–7.44
11.91
6.69
4.13
4.12 (0.77)
2.86–4.83
7.78
RUD 167 n ¼ 1
RUD 203 n ¼ 1
16.65
–
13.25
11.39
9.44
8.11
9.27
–
13.54
–
6.54
–
8.01
–
5.16
–
8.50 (0.55)
8.04–9.60
10.12 (1.48)
7.70–12.24
15.86 (1.6)
14.09–18.38
7.09 (0.77)
5.96–8.50
8.57 (1.56)
6.01–11.17
13.13 (1.64)
11.09–15.95
5.97 (0.92)
4.54–7.64
7.28 (1.24)
5.16–9.34
10.94 (1.35)
8.22–12.95
4.45 (0.55)
3.33–4.98
5.59 (0.83)
3.95–6.61
8.28 (1.04)
6.95–9.77
7.44 (0.69)
6.42–8.75
8.66 (1.42)
6.21–11.03
12.65 (1.6)
9.83–14.76
3.86 (0.38)
3.40–4.64
4.58 (0.75)
3.06–5.72
6.77 (0.76)
5.67–8.18
4.49 (0.42)
3.82–5.21
5.53 (0.86)
3.86–6.79
8.40 (1.05)
6.75–9.82
3.20 (0.36)
2.7–3.63
3.88 (0.64)
3.02–4.91
6.02 (0.67)
5.31–6.96
8.22 (0.43)
7.42–8.82
9.14 (0.56)
8.58–9.70
6.67 (0.49)
5.94–7.39
6.81 (0.53)
6.23–7.26
6.97 (0.35)
6.47–7.49
6.56 (0.84)
5.75–7.43
4.75 (0.57)
4.08–5.96
4.26 (0.80)
3.64–5.17
7.37 (0.51)
6.54–8.01
8.51 (0.45)
8.03–8.93
3.50 (0.24)
3.26–3.99
3.81 (0.65)
3.28–4.54
4.36 (0.46)
3.61–5.01
4.89 (0.51)
4.36–5.38
3.65 (0.21)
3.36–4.10
3.78 (3.54)
3.75–3.80
11.51 (1.23)
9.60–15.73
24.94 (3.16)
20.19–30.10
23.17 (1.57)
20.27–26.84
27.26 (3.28)
21.94–37.01
9.11 (0.88)
7.41–11.26
20.95 (2.52)
16.56–25.40
19.22 (2.02)
16.41–24.17
26.79 (3.46)
21.20–37.25
7.11 (0.83)
4.93–9.15
15.64 (2.13)
11.60–19.54
14.68 (1.25)
12.17–17.27
17.26 (2.53)
12.37–24.67
5.72 (0.93)
3.62–8.19
13.62 (2.1)
9.21–16.98
11.90 (1.67)
8.41–15.72
15.17 (1.96)
9.48–18.36
7.51 (2.93)
5.10–15.93
20.84 (2.73)
16.82–26.24
18.78 (1.99)
14.91–23.52
20.99 (2.35)
16.75–27.09
3.90 (0.43)
2.97–4.85
10.35 (1.81)
7.21–13.66
11.21 (0.98)
9.64–13.88
14.71 (2.11)
10.37–19.12
5.36 (0.75)
4.25–7.56
12.68 (1.87)
9.54–16.28
13.77 (1.37)
10.09–18.11
16.92 (2.12)
13.81–22.51
3.59 (0.49)
2.70–4.65
7.72 (1.43)
5.76–11.56
8.23 (0.98)
6.18–10.66
11.01 (1.33)
8.82–13.98
Other Fossils
Afropithecus n ¼ 1
Proconsul africanus n ¼ 1
Proconsul heseloni n ¼ 5
Cercopithecoid
Arboreal quadruped n ¼ 9
Semi-terrestrial n ¼ 20
Terrestrial quadruped n ¼ 10
New World monkey
Arboreal quadruped n ¼ 10
Suspensory n ¼ 4
Hominoid
Brachiator n ¼ 38
Suspensory n ¼ 32
Arboreal knuckle-walker n ¼ 52
Terrestrial knuckle-walker n ¼ 41
a
See Table 2 for variable abbreviations. For each group, mean values (with standard deviation) are listed in the first line, and the range of values is listed in the second line.
Only two variables (height and breadth of the capitate body)
could be measured on RUD 203 and together these variables do not
distinguish well amongst the different locomotor groups. Regardless, RUD 203 is virtually identical to RUD 167 in the relative size
of both of these variables and thus is likely most similar to
suspensory and brachiating hominoids and arboreal knucklewalkers (Fig. 5a, b). However, it is important to note that RUD 203
and 167 are also virtually identical to Afropithecus for both variables
Figure 2. RUD 202 scaphoid (above) in proximolateral (left) and distomedial (right) views. Comparative scaphoid morphology (below) in distomedial view. All scaphoids to 1 cm
scale except Pan and Pongo that are half scale. Oreopithecus (Basel 26) and Proconsul heseloni (KNM-RU 2036) specimens are mirror-imaged.
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
703
Figure 3. Box-and-whisker plots for each size-adjusted scaphoid variable. Box represents 25th and 75th percentiles, center line is the median, whiskers represent non-outlier
range, dots are outliers, and stars are extreme outliers. See Table 2 for variable abbreviations and Table 3 for locomotor-taxonomic group abbreviations. Oreo. ¼ Oreopithecus.
as well. Therefore, given the limited preservation of RUD 203, the
morphological similarities between RUD 203 and 167 cannot be
interpreted as strong evidence that they belong to the same taxon.
Unfortunately, the capitate is one of the few bones not known for
Epipliopithecus, a pliopithecoid like Anapithecus, and thus comparison to this taxon cannot be made. The Epipliopithecus scaphoid
shares a few similarities with RUD 202 (though morphological
comparisons can only be made to published drawings; Zapfe, 1960).
Therefore, it may be possible that both a primitive catarrhine and
primitive hominoid could share similar pleisomorphic morphology
of the capitate.
Capitate size variation
Indices of sexual dimorphism for extant taxa and Rudabánya
fossil capitates are listed in Table 5. Since only two variables could
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T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Figure 4. RUD 167 and RUD 203 capitates. RUD 167 from left to right: lateral, medial, distal, dorsal, and palmar views. In boxes, comparative capitate morphology in distal (below)
and dorsal (right) views. All capitates to 1 cm scale except Pan and Pongo that are half scale.
be measured on the RUD 203 specimen, the ISDs reported here
were calculated using the geometric mean of these two variables
only for all taxa. The geometric mean based on these two variables
was significantly correlated (rs ¼ 0.98) with the geometric mean
derived from all eight variables, and thus we feel the ISDs used here
are robust. If we assume the larger RUD 167 is male and the smaller
RUD 203 is female,2 the ISD is 1.16, which is larger than all extant
taxa except Gorilla (1.21) and Pongo (1.22).
Randomization tests revealed that the probability of sampling the
same degree of size variation from the extant sample was statistically
likely (Fig. 6) for all hominoids except P. paniscus (p ¼ 0.04). Thus, we
cannot reject the null hypothesis that RUD 167 and RUD 203 are from
a single species. Randomization tests were not run on arboreal or
terrestrial monkey groups because some are composed of different
genera or have a lower degree of sexual dimorphism than extant
hominoids and would thus be uninformative.
Discussion
Functional morphology
Qualitatively, RUD 202 has relatively distinct morphology
compared to most other Miocene hominoids that is most similar to
2
It is important to note, however, that if RUD 167 and 202 belong to the same
individual described by Kordos and Begun (1997, 2001a,b, 2002), that individual has
been described as female based on greater preservation of cranial and postcranial
anatomy. See the ‘Taxonomy’ section for more details on this sex argument.
suspensory and brachiating hominoids, particularly Pongo, and
unlike knuckle-walkers and quadrupedal monkeys in most
respects. Quantitatively, the Pongo-like morphology was not as
clear, instead showing more generalized arboreal-hominoid
morphology in some aspects. However, it is clear that the scaphoid
(and capitate) morphology is unlike terrestrial knuckle-walking
apes and more terrestrial cercopithecoids.
The distally extended lunate facet is one of the most remarkable
features of the RUD 202 scaphoid and suggests a large degree of
extension and flexion at the wrist similar to that of suspensory and
brachiating taxa. Although motion among individual carpal bones is
poorly understood for non-human primates (Orr et al., 2008),
experimental studies of humans have shown that more movement
occurs in the proximal carpal row than the distal row and particularly between the scaphoid and lunate during extension and flexion
(Garcia-Elias et al., 1991; Moritomo et al., 2006). Extant Asian apes
have a much larger range of extension (68 –85 ) and flexion (131 –
163 ) compared to African apes (extension, 29 –58 ; flexion, 117 –
135 ) and monkeys (extension, 50 –60 ; flexion, 115 –145 ; Tuttle,
1969; Richmond, 2006; unpublished data), suggesting that the
suspensory/brachiating-like lunate facet shape is also indicative of
increased mobility. However, without the adjacent scaphoid facet
on the lunate, it is difficult to make predictions of the relative degree
of mobility of this joint in Miocene hominoids.
The small, monkey-like os centrale facet may indicate a smaller
range of motion than has been found at the scaphoid-centrale
articulation in some Pongo (Orr et al., 2008). However, perhaps in
compensation for a more restricted scaphoid-centrale movement,
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
705
Figure 5. Box-and-whisker plots for each size-adjusted capitate variable. BCB and HCB only were size-adjusted by a geometric mean derived from these two variables for all
locomotor-taxonomic groups so as to include the fragmentary RUD 203 capitate (see text for details). Box represents 25th and 75th percentiles, center line is the median, whiskers
represent non-outlier range, dots are outliers, and stars are extreme outliers. See Table 2 for variable abbreviations and Table 3 for locomotor-taxonomic group abbreviations.
the os centrale facet on the proximal capitate is extensive, particularly palmarly, suggesting a larger range of midcarpal joint mobility
(see below).
Although qualitatively the shape of the RUD 202 radial facet is
more similar to that of suspensory hominoids, quantitatively it is
relatively extensive like that of quadrupedal cercopithecoids and
arboreal knuckle-walking apes. Biomechanical modelling has shown
that the majority of load from the forearm is dispersed through the
scaphoid in comparison to the lunate (Schuind et al., 1995). The
radiocarpal joint is also responsible for more than half of the range of
motion at the wrist, especially in flexion-extension and radial deviation (Crisco et al., 2005). Thus, the extensive radial facet in RUD 202
compared to other Miocene taxa may suggest (1) a relatively large
transfer of load at the radiocarpal joint, possibly signifying increased
arboreal quadrupedal activity, and/or, (2) a larger range of motion at
the radiocarpal joint.
Although the tubercle is not preserved, the neck of the scaphoid
indicates that the orientation and morphology of the tubercle is most
similar to Pongo: that is, distally extended and relatively weakly
developed compared to other African apes. A distally extended
tubercle would lengthen the carpal tunnel but would also create
a different support position for the trapezium than is found in African
apes. In Pongo, and perhaps in RUD 202 as well, the position of the
tubercle would orient the pollex in a slightly more adducted position
than that of African apes, possibly making it less efficient for
grasping. This is a reasonable functional interpretation given that the
pollex is relatively small in Pongo and not used during suspension
(Schultz, 1941; Napier, 1960; Tuttle, 1969; Jouffroy et al., 1993). This is
also consistent with the more extreme version of this tubercle
morphology in Oreopithecus, which is described as having a thumb
that is ‘‘quite short’’ (Harrison, 1991: 289; contra Moyà-Solà et al.,
1999, 2005; Susman, 2004). The RUD 202 scaphoid could similarly be
associated with a more adducted, and possibly reduced, pollex that is
often associated with suspensory behavior. However, the fossil
evidence to support this hypothesis has yet to be recovered.
The RUD 167 capitate qualitatively appears more generalized in
its functional morphology than that of the scaphoid. However,
quantitatively, it has a relatively strong suspensory and brachiating
hominoid-like functional signal. RUD 167 is best described as having
a generalized arboreal morphology with indications of increased
mobility. In particular, the palmarly expanded capitate-centrale
articulation suggests an increase in radial deviation at the midcarpal
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T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
Figure 5. (continued).
joint. In radial deviation, the scaphoid undergoes palmar flexion
(Linscheid, 1986; Zylstra, 1999) so the palmar extension of the
capitate’s proximal facet for the os centrale may imply a greater
potential for radial deviation. Radial deviation is typical in climbing
and suspending hand postures (Tuttle, 1969; Sarmiento, 1988).
Hylobatids, which have a similar palmar extension of the proximal
capitate facet, have a larger range of radial deviation than African
apes (Tuttle, 1969), which have less extensive proximal facets.
Although the hamate facet also shows similarities to suspensory
and brachiating hominoids, previous analyses (Kivell, 2007) have
shown that the shape of the hamate facet carries little functional
significance given the relatively immobile articulation shared
between the capitate and hamate (Garcia-Elias et al., 1991). The
second and third metacarpal facets are hominoid-like and suggest
increased stability in a larger range of motion, including adduction
and abduction of digits, than is found in monkeys. The mediolaterally narrow capitate head and distal capitate body suggest
that they may experience relatively weaker compressive loading
than that of knuckle-walking apes and quadrupedal cercopithecoids. The fact that RUD 167 looks almost identical to that of
Proconsul or Dryopithecus (Moyà-Solà et al., 2004) in the overall
shape of the body suggests that the capitate largely retains the
plesiomorphic morphology. Similarities to suspensory and brachiating hominoids suggest that these extant taxa, at least in the
manner in which the morphology is broadly quantified here, also
retain relatively plesiomorphic morphology (although qualitatively
the morphology looks more derived and brings to light the value of
using more detailed, quantitative measures in future analyses).
RUD 167 is best described as having the morphology of a generalized arboreal hominoid with modifications for increased mobility
consistent with some suspensory and climbing capabilities.
The second and third metacarpal facets preserved on the RUD
203 capitate are qualitatively very similar to those of RUD 167
described above. Although RUD 203 comes from an individual
about half the size of RUD 167, quantitatively the morphology is
virtually identical when adjusted for this size difference. However,
because the same is true of Afropithecus, it cannot be assumed that
the remainder of the RUD 203 capitate would be equally similar to
that of RUD 167 and functionally indicative of the same generalized,
Table 5
Indices of sexual dimorphism (ISD; mean male/mean female geometric mean
values) in extant hominoidsa
Species
_n
_ mean
RUD 167 and 203
Gorilla
Pan paniscus
Pan troglodytes
Pongo
Hylobatids
1
17
9
12
11
8
11.18
23.18
16.27
18.07
20.22
8.79
a
S.D. ¼ standard deviation.
_ S.D.
\n
\ mean
\ S.D.
ISD
1.53
0.84
1.27
1.72
0.39
1
17
9
13
16
7
9.61
19.13
15.37
16.61
16.59
8.31
1.37
0.69
1.33
0.92
1.01
1.16
1.21
1.06
1.09
1.22
1.06
T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
707
Figure 6. Frequency and probabilities (%) of size ratios in all possible pairwise comparisons of the capitate plotted for Pongo, Gorilla, Pan paniscus, and Pan troglodytes. The size ratio
of RUD 167/RUD 203 (1.16) is marked by the dotted line. (*) Probability less than 5%.
arboreal hominoid-like morphology with increased mobility for
suspension and climbing activities.
Taxonomy
Both the RUD 202 scaphoid and RUD 167 capitate are of
a compatible size and share similar functional morphology. The
functional interpretation of both as a generalized, arboreal suspensory primate is consistent with the interpretations of other postcranial material, including phalanges, distal humerus, and femur, of
Rudapithecus (Begun, 1988, 1992, 1993). Although Anapithecus is
considered to have engaged in some suspensory locomotion as well
(Begun, 1988, 1993), carpal size suggests that it is of a similar body
size as that of extant Papio and outside the estimated body mass
range of Anapithecus (Begun and Kordos, 2004). Thus, both RUD 202
and RUD 167 are confidently classified as Rudapithecus hungaricus
and are possibly from the same individual. Since the discovery of
these carpal bones, numerous other associated cranial, dental, and
postcranial remains have been uncovered at Rudabánya that appear
to be from the same, small female Rudapithecus (Kordos and Begun,
1997, 2001a,b, 2002). The scaphoid and capitate are functionally and
morphologically consistent with these other fossils and may well be
part of the same Rudapithecus individual.
The RUD 203 capitate is more difficult to assess taxonomically.
The preserved morphology is remarkably similar to that of RUD 167,
indicating that it is a smaller Rudapithecus individual. However, its
size suggests that it is of a primate about half the size of that RUD
167, which is closer to that of Anapithecus. The largest body mass
estimates for Rudapithecus come from the distal humerus, which is
the size of a male chimpanzee, roughly 40 kg (Morbeck, 1983;
Begun, 1994). Although a range of body mass between roughly
10–40 kg for Rudapithecus would be extreme within a single taxon,
the degree of sexual dimorphism derived from the geometric mean
(ISD) between the fossil pair is lower than that of Gorilla and Pongo,
suggesting that this amount of size variation is not unreasonable.
Further, randomization tests revealed that it was statistically likely
to find two individuals with this degree of size variation in all
hominoids except P. paniscus. These results must be considered
with caution as they are based on a geometric mean of carpal
variables and the carpal bones are not likely to be highly correlated
with true body size. Nevertheless, the data presented here, both
morphological and quantitative, indicate that the single-species
hypothesis for RUD 203 and RUD 167 cannot be falsified. However,
this conclusion has important implications. If the RUD 202 scaphoid
and RUD 167 capitate are both considered to be associated with the
same small female individual, as is supported by this analysis, than
RUD 203 must represent a third, even smaller, size category. This is
a large and unlikely range of size variation within one sex. Therefore, in the end, given the small number of comparative measurements possible between the two RUD capitates, the similarities to
other fossil catarrhines, the large difference in size, and the absence
of data on pliopithecoids including Anapithecus, we consider the
issue of the taxonomic status of RUD 203 unresolved.
Conclusions
We attribute two of the first primate carpal bones (RUD 167
and 202) described from Rudabánya to the hominid Rudapithecus
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T.L. Kivell, D.R. Begun / Journal of Human Evolution 57 (2009) 697–709
hungaricus. A third, fragmentary specimen (RUD 203) from a much
smaller individual cannot be assigned to a taxon. Qualitative and
quantitative analyses indicate that the functional morphology of
the scaphoid and capitates is hominoid-like and consistent with
arboreal locomotion, including more suspensory and climbing
activities than is typical of arboreal or terrestrial monkeys. The
scaphoid is most similar to Pongo in its morphology and indicates
a large degree of mobility. We interpret the morphology of the
Rudapithecus capitate as preserving a more plesiomorphic hominoid condition, similar to that of Proconsul or Dryopithecus, that is
best described as a generalized arboreal hominid with indications
of enhanced mobility at the midcarpal joint. The poorly preserved
smaller capitate cannot be distinguished functionally from the
larger specimen. This functional interpretation is consistent with
that of other postcranial remains from this taxon (Begun, 1988,
1992, 1993, 1994) and further supports the conclusion that
Rudapithecus is an arboreal ape, capable of more suspension,
climbing, and quadrupedalism than the early Miocene hominoids
but lacking all the distinct locomotor specializations of any one
extant hominoid taxon.
Acknowledgements
We are grateful to László Kordos for providing access to the
Rudabánya fossil material and to Mike Rose and Alan Walker for
providing the Miocene fossil casts. We thank the following curators
and institutions for access to extant specimens: W. Wendelen and
M. Louette (Musée Royal de l’Afrique centrale); M. Harman (PowellCotton Museum); L. Gordon (Smithsonian Institution); Y. HaileSelassie and L. Jellema (Cleveland Museum of Natural History);
J. Sirianni and E. Hammerl (SUNY University at Buffalo); J. Eger and
S. Woodward (Royal Ontario Museum); F. Burton (University of
Toronto). We also thank the anonymous reviewers whose
constructive comments greatly improved this paper. This research
was funded by the Natural Sciences and Engineering Research
Council (NSERC) of Canada to both authors, an Alexander von
Humboldt Stiftung Research Fellowship to DRB, and General
Motors Women in Science and Mathematics Award and University
of Toronto Travel Grant to TLK.
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