THE ANATOMICAL RECORD 294:217230 (2011)

Geometric Morphometric Analysis of

Tibial Shape and Presentation Among
Catarrhine Taxa
KEVIN TURLEY,* EMILY H. GUTHRIE, AND STEPHEN R. FROST
Department of Anthropology, University of Oregon, Eugene, Oregon

ABSTRACT
The proximal component of the talo-crural joint, the tibia, was com-
pared, using geometric morphometrics, in 240 specimens from 10 extant
taxa to identify differences in shape and the factors inuencing them.
The specimens were laser scanned, digitally reconstructed, and land-
marked. Regression analysis was used to evaluate tibial shape, and sig-
nicant amounts of shape variation among taxa were due to body mass,
tibial size, superfamily, and substrate preference in the whole tibia, as
well as, separate analysis of the distal tibial articular facets, and the
medial malleolar facet. The most important factor for whole tibial shape
was tibial robusticity, which closely correlated with body mass. However,
substrate preference was also a signicant factor in tibial shape and inde-
pendent from body mass. Substrate preference was also the most impor-
tant factor dening distal articular morphology. Principal components
analysis and pairwise permutation tests were used to compare differences
in morphology among taxa. Nearly all were signicantly different in over-
all tibial shape, and distal morphology. Shape and presentational mor-
phology associated with body mass, tibial size, superfamily, and substrate
preference were identied, along with the similarities and differences
among individual taxa. These were visualized by TPS deformation of an
exemplar surface. Relationships among these factors were assessed with
their dot-product. Results demonstrated that size signicantly inuenced
proximal presentation, while substrate preference inuenced articular
morphology. Anat Rec, 294:217230, 2011. V C 2010 Wiley-Liss, Inc.

Key words: tibial shape; substrate preference; ankle

homoplasty

INTRODUCTION
The talo-crural joint is a highly conserved feature
among Primates (Vancata, 1991; Hall, 1998; Lieberman
et al., 2001; Wagner, 2001). It is a structural unit that
forms the interface between the leg (and the rest of the or-
ganism) and the foot (and substrate) (Lewis, 1989; Ruff,
2002). It can be considered to consist of three elements;
proximal presentation, appositional articular morphology,
and distal presentation. Presentation is the orientation of
the joint surfaces in space relative to both the organism
and substrate. It differs from alignment in being inde-
pendent of other structures, other than inuencing their
potential shape. It is a function of both proximal and dis-
tal bone shape, and constrains the variation observed in
articular morphology. For the talo-crural joint, proximal
presentation consists of tibial shape, the tibiotalar axis
and plane, and the orientation of the distal articular fac-
ets. Distal presentation consists of talar shape, articular
facet shape, and angulation. Appositional articular mor-
phology is the shape of the joint surfaces themselves
which provide the range of joint function for the organism
constrained by both proximal and distal bone shape.
Grant sponsor: NSF; Grant numbers: BCS-0452538, IIS-
513660; Grant sponsor: University of Oregon.
*Correspondence to: Kevin Turley, Department of Anthropol-
ogy, University of Oregon Eugene, Oregon 97403-1218. Tel.:
541-346-5161, Fax: 541-346-0668. E-mail: kturley@uoregon.edu
Received 28 May 2010; Accepted 29 September 2010
DOI 10.1002/ar.21307
Published online 3 December 2010 in Wiley Online Library
(wileyonlinelibrary.com).

V
C 2010 WILEY-LISS, INC.
218 TURLEY ET AL.

TABLE 1. Number of specimens of different taxa used in this analysis

Taxon N M F Mass (kg) M/F Substrate (010) M/F
Homo sapiens 63 30 31 59.3/53.2 9.99/9.99
Pan troglodytes 63 29 31 56.6/44.0 4/3
Gorilla gorilla 45 26 14 169.8/73.5 6/4
Pongo sp. 15 9 5 78.1/35.7 6/0.2
Hylobatidae 6 1 4 8.6/8.0 0.1/0.1
Macaca fascicularis 8 4 3 5.3/3.6 1/1
Macaca thibetana 9 7 1 15.2/9.5 8/8
Papio hamadryas 12 5 5 25.1/13.3 9/9
Nasalis larvatus 10 8 2 20.4/9.8 0.1/0.1
Colobus guereza 8 4 3 13.5/9.2 0.1/0.1
Mass values from Smith and Jungers (1997) and Delson et al. (2000). For several taxa mass values pooled as weighted
averages from subtaxa as follows: G. gorilla derived from two subspecies, G. g. beringei (three specimens) and G. g. gorilla
(36); H. sapiens six populations (see text); Hylobatid specimens from four species masses pooled Hylobates hoolock (1), H.
lar (1), H. muelleri (1), S. syndactylus (3); Pan troglodytes from three subspecies P. t. schweinfurthii (8), P. t. troglodytes
(50), P. t. verus (4); and Pongo from two species P. pygmaeus (6) and P. abelii (9). Papio hamadryas specimens are all P. h.
anubis. Substrate use from Fleagle (1999).

Fig. 1. Thirty-one landmarks on the tibia used in this study on the articular surfaces and tibial tuberos-
ity illustrated using a tibia of a male Gorilla gorilla.

Variation in presentation and articular morphology
reect natural selection for its genetically canalized
state and the response of the organism to the biome-
chanical stresses encountered during ontogeny. Both of
these factors produce the endpoint morphology encoun-
tered in the individual (Lovejoy et al., 2000; Lieberman
et al., 2001; Pearson, 2004; Hall, 2005). Variation in this
endpoint morphology allows for the differences in func-
tional morphology of this joint complex among taxa. As a
result, talo-crural morphology reects the different phy-
logenetic histories, modes of locomotion, and substrate
preferences observed among primates.
In this study, we use 3-D geometric morphometric
methods to examine differences in shape of the proximal
element of the talo-crural joint, the tibia, among catar-
rhines. In particular, we examine the relationship of
multiple factors that relate to the biological role of the
talo-crural joint (sensu Bock and von Wahlert, 1965).
These factors have been previously demonstrated to be
related to variation in shape of articular morphology,
and include taxon, body mass, tibial size and robusticity,
and substrate preference (Latimer et al., 1987; Jungers
1988; Lieberman et al., 2001; Aiello and Dean, 2002;
Ruff, 2002; Pearson, 2004; Gebo and Schwartz, 2006;
DeSilva, 2009). We also examined differences and simi-
larities in mean tibial shape of the diverse Catarrhine
taxa that formed the study group.
MATERIALS AND METHODS
Sample
The tibiae of 240 specimens from 10 Catarrhine taxa
formed the study sample. All were adult, without pathol-
ogy, and among the nonhuman specimens, wild-shot
 DETERMINANTS OF TIBIAL SHAPE 219
TABLE 2. Landmarks used in this study Smith and Jungers, 1997; Katzmarzyk and Leonard,
1998; Fleagle, 1999; Delson et al., 2000). Substrate pref-
Number Type Description
erence was estimated from 0 (most arboreal) to 10 (most
1 II Most anterior point on the medial facet terrestrial). For higher taxa, mass and substrate values
2 III Antero-medial prominence (anterior were averaged and weighted by the number of individu-
cruciate ligament insertion) of the als in each sub-taxon (Table 1).
medial facet
3 III Postero-medial prominence of
the medial facet Data Collection
4 III Postero-medial inection of the
medial facet Laser surface scans were made from each specimen
5 II Most posterior point on the medial facet using a Konica Minolta Vivid 910 Noncontact 3-D Digit-
6 III Postero-lateral inection of the izer, and were processed using Geomagic Studio 8 soft-
medial facet ware. Thirty-one landmarks were collected by one
7 II Most medial point on the medial facet observer (KT) using Landmark Editor software (Wiley,
8 III Antero-lateral inection of the 2006). These covered the proximal and distal articular
medial facet surfaces as well as the tibial tuberosity (White and Folk-
9 III Center of medial facet concavity ens, 2000) (Fig. 1, Table 2). Ten were Type II landmarks
10 III Tibial tuberosity
11 II Most anterior point on the lateral facet and 21 were Type III landmarks chosen to reect both
12 III Antero-lateral inection of the function and the overall size of the bone (Bookstein,
lateral facet 1991; Harcourt-Smith, 2002). Three landmark subsets
13 II Most lateral point on the lateral facet were used: the whole tibia, distal tibial facets (trochlear
14 III Posterior point of inection of the and medial malleolar), and medial malleolar facet alone.
lateral facet The proximal tibia and trochlear facet alone were also
15 II Most posterior point on the lateral facet considered, but did not yield any signicant shape differ-
16 III Postero-medial prominence of the ences among taxa or related to the other variables con-
lateral facet sidered in this study, and so are were not considered
17 III Antero-medial prominence of the
lateral facet further. Separate analyses were performed on each sub-
18 III Antero-medial inection of the set to determine shape differences in each region with-
lateral facet out the confounding effects of the whole tibial
19 III Center of the lateral facet concavity morphology.
20 II Most postero-lateral point on the To evaluate observational error, one specimen of Homo
trochlear facet sapiens was landmarked ten times. The 93 individual
21 III Most medio-lateral point (midpoint) landmark coordinates demonstrated an average stand-
on the trochlear facet ard deviation of 0.5 mm (range 01.7 mm, with 89 coor-
22 II Most antero-lateral point on the dinates <1.0 mm). The distance from the centroid to
trochlear facet
23 III Most anterior point (midpoint) on the each of the 31 landmarks had a standard deviation of
trochlear facet 0.547 mm (range 0.2220.989 mm). Principal Compo-
24 III Most antero-superior point on the nents Analysis revealed tight clustering of these
medial malleolar facet repeated measures compared to variation within and
25 III Most superior point on the medial among taxa used in this analysis. Since this study
malleolar facet focused on variation above the species level, precision
26 III Most postero-superior point on the was deemed satisfactory.
medial malleolar facet
27 III Most posterior point (midpoint) on
the trochlear facet Generalized Procrustes Analysis
28 III Center of the trochlear facet
29 II Anterior trochlear facet medial Generalized Procrustes Analysis (GPA) was performed
malleolar junction using Morpheus (Slice, 1998). GPA superimposes land-
30 III Center trochlear facet medial mark congurations and removes variance due to posi-
malleolar junction tion and rotation, and scales each to unit centroid size
31 II Posterior trochlear facet medial (Rohlf and Slice, 1990). Centroid size is the square root
malleolar junction of the sum of the squared distances of each landmark to
the centroid (Rohlf and Slice, 1990), and is stored as a
separate variable during GPA. As our data set included
with provenience documented. The 63 human specimens specimens from both the right and left sides, GPA was
derived from six populations: 10 European American done with reection allowed. Separate GPAs were per-
(19th Century), 10 African American (19th Century), 10 formed for each of the three landmark subsets. All sub-
Inuit (19th Century), 10 Egyptian (4th Century), 10 sequent statistical analyses were performed using SAS
Southwest Paleoamericans, and 13 California Paleoa- 9.1 (SAS Institute, Cary, NC). As GPA aligned coordi-
mericans. Nonhuman primate specimens included 129 nates have a very high correspondence with their Eu-
Apes and 48 Old World monkeys (see acknowledgements clidean tangent space projections, un-projected aligned
for specic institutions) (Table 1). The latter were chosen coordinates were used. Shape differences among land-
to represent both subfamilies (Xing et al., 2007) with a mark congurations were measured by Procrustes dis-
range in size and substrate use to the extent possible. tance; the Pythagorean distance between the two
Estimated mass and substrate for all taxa were obtained Procrustes superimposed landmark congurations
from the literature (Kraus, 1961; Auger et al., 1980; (Bookstein, 1991). Procrustes distances among taxon
220 TURLEY ET AL.

means were compared using minimum spanning trees
for each subset in NTSYSpc. 2.2 (Rohlf, 1998).
Principal Component Analysis
Principal component analysis (PCA) was performed on
the covariance matrix of the GPA superimposed land-
mark coordinates as a data reduction and exploration
TABLE 3. Aspects of tibial shape examined in
each data set
Whole tibia data set

Robusticity

Medial angulation

Axial rotation (tibial torsion) (Aiello and Dean, 2002)

Plane (medial to lateral angulation of the tibiotalar
joint surface ) (Aiello and Dean, 2002), angulation
increases as the lateral border moves superiorly

Axis (posterior to anterior angulation of the
tibiotalar joint surface) (Aiello and Dean, 2002)
angulation increases as the anterior border
moves superiorly
Distal tibial facets data set

Tibial shape-oval/trapezoid

Anterior trochlear margin (plane, lateral to
medial) distinct from the whole tibia angulation
increases with superior displacement of the
antero medial margin

Posterior trochlear margin (plane, lateral to medial)
distinct from the whole tibia angulation increases
with superior displacement of the postero medial
margin

Medial trochlear margin (trochlear medial malleolar
groove) (axis posterior to anterior)

Lateral trochlear margin (axis posterior to anterior)

Trochlear-medial malleolar facets angle

Central trochlear concavity (depth)

Relative trochlear-medial malleolar facets surface
area ratio
Medial malleolar facet data set

Relative height (midline)

Relative length (base)

Base shape (concave to convex)

Central shape (concave to convex)

Apex shape (position, posterior-central-anterior)
technique (Sokal and Rohlf, 1995). Initial visualization
of the effect on shape described by each Principal Com-
ponent (PC) both within and among superfamilies was
performed using Morphologika software (OHiggins and
Jones, 1998; OHiggins and Jones, 2006). Scores for PC
1-5 were compared between taxa based on phylogeny,
body mass, geographic distribution, and substrate pref-
erence, with results reported with a Bonferroni-adjusted
signicance level of P < 0.001.
Permutation Test
To test for differences in shape between individual
taxa, pair-wise permutation tests with 1000 replicates
were performed for each landmark subset. Individuals
were randomly permuted across the two taxa and Pro-
crustes distance was calculated between the permuted
groups means. The a was the fraction of permuted val-
ues that were greater than the actual Procrustes dis-
tance between group means (Good, 2000). Results were
reported with a Bonferroni-adjusted signicance level of
P < 0.001.
Regression Analysis
Multivariate regression analysis was performed to
assess differences in shape related to different variables
(Bookstein, 1996; Frost, et al., 2003). GPA aligned coor-
dinates were used as dependent variables with substrate
preference, body mass, tibial size (as measured by log
transformed centroid size), superfamily (Hominoidea and
Cercopithecoidea), and tibial robusticity (as measured by
a tibial index of maximum breath/maximum length)
used as independent variables.
The proportion of the total variance accounted for by
each independent variable (robusticity, mass, log cent-
roid size, phylogeny, and substrate preference) was cal-
culated for each of the subsets by subtracting the
residual variance after regression from the total var-
iance and dividing the difference by the total variance
(multivariate multiple regressions) (Frost et al., 2003).
To evaluate the relationship among the independent var-
iables, the angles among the shape vectors for each

TABLE 4. The rst ve eigenvalues of the covariance matrix on Procrustes aligned coordinates for
Principal component analyses on the three different data sets
Rank Eigenvalue Difference Proportion Proportion
Whole tibia
1 0.00074234 0.00056530 0.5364 0.5364
2 0.00017704 0.00009602 0.1279 0.6643
3 0.00008102 0.00002964 0.0585 0.7229
4 0.00005138 0.00001049 0.0371 0.7600
5 0.00004089 0.00000927 0.0295 0.7895
Distal tibia
1 0.00324367 0.00057662 0.1634 0.1634
2 0.00266705 0.00084367 0.1344 0.2978
3 0.00182338 0.00018250 0.0919 0.3896
4 0.00164088 0.00031165 0.0827 0.4723
5 0.00132923 0.00023975 0.0670 0.5393
Medial malleolar facet
1 0.00699940 0.00045936 0.2201 0.2201
2 0.00654004 0.00192740 0.2057 0.4258
3 0.00461264 0.00162124 0.1451 0.5709
4 0.00299140 0.00020189 0.0941 0.6650
5 0.00278951 0.00062447 0.0877 0.7527
 DETERMINANTS OF TIBIAL SHAPE
Fig. 2. Scatter plot showing principal component scores for individual tibiae based on PCA of Pro-
crustes aligned coordinates. PC1 is on the X-axis and PC 2 on the Y-axis. Convex polygons are used to
show the range of scatter with individual points hidden for clarity.
factor were calculated as the arccosine of their vector
correlation (dot product) (Cobb and OHiggins, 2004).
Parallel angulation (0 ) demonstrated correlation, or-
thogonal angulation (90 ) demonstrated independence.
Visualization
Visualization of shape differences was accomplished by
comparing landmark congurations directly in Morpheus
and by warping an exemplar surface to t those land-
mark congurations using Landmark Editor. These were
done separately for each of the landmark subsets. Mean
landmark congurations were computed for each taxon.
Shape differences associated with substrate preference,
mass, log tibial size, superfamily, and, for the whole
tibia, tibial index were visualized by adding the vector of
regression coefcients from multivariate regression to
the consensus landmark conguration for each of the
landmark subsets. Features used in the resulting
descriptions are explained in Table 3.
RESULTS
Principal Component Analysis
The eigenvalues, difference, proportion, and cumula-
tive percentage for each of the rst ve PCs for each
landmark subset are presented in Table 4. When scores
from the rst ve PCs are plotted, clear shape differen-
ces among taxa are evident by the separation among
their constituent individuals (Fig. 2). Tibial robusticity
correlated highly (r 0.98) with PC-1 of the whole tibial
data set. The rst ve PCs constituted 79% of the var-
iance in the whole tibia, 54% in the distal tibial facets,
and 75% in the medial malleolar facet (Table 5).
Analysis of PC 1-5 of selected taxa revealed Gorilla
and Pan differed in all ve in the whole tibia and distal
221
tibial facets, and three in the medial malleolar facet (PC
1,3,4) (P < 0.05) Homo and Pan differed in all ve in the
whole tibia, and four in the distal tibial facets (PC
1,2,4,5), and medial malleolar facet (PC 1,2,3,4) (P <
0.05) but Gorilla and Homo differed in only two in the
whole tibia (PC 1,2), distal tibial facets (PC 2,3) and
medial malleolar facet (PC 1,3) (P < 0.05). Among the
cercopithecoids examined, Papio and Macaca thibetana
differ in only two in all three data sets (PC1, 3; 1,2; and
2,3) (P < 0.05) while Colobus and Nasalis differed in
only one in the whole tibia, and distal tibial facet subset
(PC 3; 2) (P < 0.05). Of note, phyolgenetic comparative
methods were not used in these comparisons, and thus,
the statistical signicance of inferences needs to be
explored (Rohlf, 2006).
Differences in Shape Related to Biological Role
The effect of each factor studied (substrate preference,
body mass, log of tibial size, superfamily, and tibial
index for the whole tibia) for shape in each landmark
subset was highly signicant (P < 0.0001). For the whole
tibial data set, the factors that accounted for the most
variance were robusticity, mass, and superfamily, which
represented 51.5%, 30.3%, and 18.2% of total variance
respectively. These factors were closely related, with an
angle of only 7.5 , almost parallel, for robusticity and
mass, while mass and superfamily were 26.2 , and
robusticity and superfamily 30.5 . Substrate preference,
on the other hand accounted for only 10%, but was or-
thogonal from all other factors (robusticity74.2 ,
mass80.2 , superfamily79.8 ), excepting log tibial
size (38.3 ), which may reect the effect of the Homo
sample. For the distal tibial and medial malleolar data
sets, however, substrate preference accounted for 6.1%
222 TURLEY ET AL.

TABLE 5. Description of shape differences associated with the rst ve principal components for each
data set
Whole tibia data set
PC-1 reects tibial robusticity, from gracile to robust (with tibial index highly correlated (r 0.98), while length relative to
centroid size (r 0.42) and length (N.S.) are not, and medial angulation of the distal tibia increasing proximal platform
presentation in robust taxa to the arboreal substrate.
PC-2 reects axial rotation of the tibia medially and laterally (tibial torsion).
PC-3 reects distal shape change (from decrease antero-posterior (AP) and increased medio-lateral (ML) to increased AP
and decreased ML, and oval to trapezoid), and plane change from anterior to posterior.
PC-4 shows proximal facet rotation, narrowing of the trochlear facet, and decrease in the trochlear-medial malleolar angle.
PC-5 shows narrowing of the posterior trochlear facet, angulation of the trochlear facet axis.
Distal tibia data set
PC-1 reects change in AP-ML dimensions from oval to trapezoid, decrease of the malleolar facet area relative to the troch-
lear facet area, and medial malleolar base length, and increase in the medial malleolar convexity.
PC-2reects trochlear surface change from curved to at and anterior displacement of the medial groove from symmetrical.
PC-3 reects a change in medial groove and facet shape from anteriorly displaced to central
PC-4 reveals a change in trochlear malleolar angulation with decreased angle and concavity to at.
PC-5 narrowing of the posterior trochlear facet width, and angulation of the trochlear facet axis
Medial malleolar facets data set
PC-1 revealed a change from low and at to high central apex.
PC-2 revealed a change from at base with apex anterior to centrally elevated base with central apex.
PC-3 revealed a change from a concave base to a convex base.
PC-4 revealed a change from a concave to convex facet and apical displacement from anterior to posterior.
PC-5 revealed a change in the central base from concave to convex.

TABLE 6. Percent of total variance explained by different factors within different landmark subsets
Data set Substrate Robusticity Mass Log centroid size SuperFamily
Whole tibia 10.6% 51.5% 30.3% 8.1% 18.2%
Distal tibia 6.1% 6.8% 4.7% 4.3% 3.1%
Medial malleolus 6.6% 3.4% 2.5% 6.0% 2.9%

and 6.6% of the total variance, again independent of
other factors including log tibial size. Mass and super-
family accounted for less in each of these subsets (4.7%
and 3.1% in the distal tibia, and 2.5% and 2.9% in the
medial malleolus) with an interrelationship demon-
strated in each. The log of tibial size accounted for 4.3%
of the distal tibial, and 6.0% of the medial malleolar var-
iance, and was independent of substrate in both, and
robusticity in the medial malleolar subset. Robusticity
accounted for 6.8% and 3.4%, respectively, and was
interrelated to all factors except substrate in both
groups and log tibial size in the medial malleolar subset
(Tables 6 and 7). Shape differences associated with each
of these factors were examined in Morpheus and Land-
mark Editor and are summarized below (Fig. 3).
medial malleolar facet of more arboreal taxa was longer
with a central apex and a convex base.
Body mass and robusticity. The whole tibia was
associated with symmetrical widening of the proximal
facets, the trochlear facet, and medial rotation of the dis-
tal tibia. There was increased lateral to medial plane of
the tibiotalar joint surface, medial angulation and
medial rotation of the distal tibia. The distal tibial facets
were enlarged with widening of the angle between the
trochlear and medial malleolar facets. In taxa with more
robust tibiae, the medial malleolus was enlarged in
width and height and had less convexity of the facet,
and in taxa of greater mass it had less convexity, with
central elevation of the base and an anterior displace-
ment of the apex.

Substrate. The whole tibia of more terrestrial taxa
has more lateral rotation (torsion) of the distal end and
less medial angulation (a nding unique to Gorilla, Pan,
and Pongo-angulation of the distal 1/3-absent in Homo,
Hylobates, and all Cercopithecoids) than do more arbo-
real forms. The distal tibia of more terrestrial taxa also
showed a atter lateral to medial plane of the tibiotalar
joint surface (Aiello and Dean, 2002) and a less oval,
more trapezoidal trochlear facet with a deeper central
concavity and atter anterior margin. The distal tibiae
of more arboreal taxa had anterior, medial and superior
displacement of the anterior margin and trochlear-
medial malleolar groove, with oval lateral and posterior
margins. The medial malleolar facet of more terrestrial
taxa was low and at with a concave base, while the
Tibial size. In larger tibiae (as measured by log cent-
roid size), wider proximal, and trochlear facets were
observed, along with lateral rotation of the distal end.
The distal tibial facets of larger tibia showed an enlarge-
ment of the medial malleolar facet relative to the troch-
lear facet, and a change in shape from rectangular to
triangular. The medial malleolar facet of larger tibia
showed less convexity of the malleolar facet with the
apex moving anterior.
Superfamily. Superfamily showed, in the whole
tibia, wider proximal and trochlear facets in hominoids
than cercopithecoids. In the distal tibial facets, there
was a decrease in an antero-posterior dimension and
increase in medio-lateral dimension of the trochlear fact
 DETERMINANTS OF TIBIAL SHAPE 223
TABLE 7. Angular differences (dot product) between the vectors for different factors, presented in
degrees
Substrate Robusticity Mass Log size Superfamily
Whole tibia
Substrate 105.8 99.8 38.3 79.8
Robusticity 74.2 7.5 70.2 30.5
Mass 80.2 7.5 64.3 26.3
Log size 38.3 70.2 64.3 42.9
Superfamily 79.8 30.5 26.3 42.9
Distal tibia
Substrate 117.7 74.1 55.7 92
Robusticity 62.3 31.2 55.9 47.8
Mass 74.1 31.2 30.2 46.2
Log size 55.7 55.9 30.2 53.6
Superfamily 88 47.8 46.2 53.6
Medial malleolus
Substrate 119.1 92.4 66.2 86.1
Robusticity 60.9 48.4 68.1 46.4
Mass 87.6 48.4 35.0 45.5
Log size 66.2 68.1 35.0 36.6
Superfamily 86.1 46.4 45.5 36.6

from cercopithecoids to hominoids. Finally, there was a
decrease in the surface area of the medial malleolar
facet relative to the trochlear facet from cercopithecoids
to hominoids.
Differences in Shape Among Taxa
The magnitude of differences in shape between species
means as measured by Procrustes distance for the whole
tibial data set, distal articular facet, and medial malleo-
lus are presented in Table 8. These illustrate that shape
is highly conserved within taxa, but there appears to be
an inuence of multiple factors on tibial shape, including
robusticity and presentation in the whole tibial data set
and substrate preference in the articular morphology.
The results of the pair-wise permutation tests for dif-
ferences in mean shape between different taxa are pre-
sented in Table 8. Signicant differences among taxa
were found in the whole tibial data set between all pairs
except Colobus versus Nasalis and Macaca fascicularis
versus Nasalis, even after applying the Bonferroni cor-
rection. The distal tibial facet was also different for all
comparisons of apes versus monkeys. Within hominoids
all were different except for Pongo versus Hylobates and
among cercopithecoids all taxa showed differences
except, Macaca fascicularis versus M. thibetana, M. thi-
betana versus Nasalis, and Nasalis versus Colobus.
Finally, for comparisons of the medial malleolar facet set
Homo was different from all taxa; Pan was different
from all taxa except Nasalis; Gorilla was different from
all taxa except Macaca thibetana; Pongo was different
from all taxa except Hylobates and Nasalis; Papio was
different from all taxa except Macaca thibetana and
Colobus. No signicant differences were noted among
the remaining taxa. The mean shapes for each taxon
were examined in Morpheus and Landmark Editor and
are summarized below (Table 3). These differences in
the distal tibial facets and the medial malleolar facet are
illustrated in four selected taxa, Papio, Pan, Gorilla,
and Homo (Figs. 4, 5).
Gorilla spp. demonstrated the greatest robusticity and
medial angulation of the distal 1/3 of the tibia. They
have moderately elevated plane and axis of the tibiotalar
joint surface, and moderate medial axial rotation accen-
tuated by the effects of presentation (angulation, plane
and axis). Their trochlear shape was trapezoidal with a
marked wedge shape. They had slightly convex anterior,
posterior, and lateral margins, and a concave medial
margin. The anterior plane was mildly elevated from lat-
eral to midpoint then sharply increased from the mid-
point to the antero-medial margin and displaced
superiorly. The posterior plane was concave and slightly
increased lateral to medial. The lateral margin axis was
increased posteriorly to anteriorly, and the medial mar-
gin was convex with the axis markedly elevated and
superiorly displaced from midpoint to the anterior
medial point. The trochlear-medial malleolar angle was
increased. The central concavity was mild and trochlear-
medial malleolar facet area ratio was 3:1. The medial
malleolar relative height was moderate and length
increased, and its base was mildly convex as was the
central shape with the apex anteriorly displaced.
Pan troglodytes demonstrated decreased robusticity
and medial angulation of the distal 1/3 of the tibia com-
pared with Gorilla. There was a more marked elevation
of plane but decreased axis of the tibiotalar joint surface,
and less medial axial rotation. The trochlear shape was
oval with a slightly concave anterior and medial margin,
and the anterior plane was markedly elevated and supe-
riorly displaced from the antero-lateral to antero-medial
margin. The posterior plane was concave and slightly
increased lateral to medial. The lateral axis was
decreased, while the medial axis was convex and mark-
edly increased from the midpoint to antero medial cor-
ner. The trochlear-medial malleolar angle was decreased
(in the mid-malleolar medial projection). The central
concavity was moderate and trochlear-medial malleolar
facet area ratio was 3:1. The medial malleolar relative
height was increased, relative length decreased from Go-
rilla, and the base was more convex and the central
shape more convex, with the apex central.
224 TURLEY ET AL.
Fig. 3. Visualization for the whole tibial data set of four primary var-
iables analyzed: log of centroid size (A), body mass (B), substrate pref-
erence (C), and superfamily (D). Landmark congurations estimated by
adding and subtracting regression coefcients to the concensus con-
Homo sapiens demonstrated decreased robusticity
with absence of medial angulation of the distal 1/3 of the
tibia. There was a at plane and moderately elevated
axis with lateral axial rotation of the tibiotalar joint sur-
face. The trochlear shape was trapezoidal but less wedge
guration, these were magnied for the purpose of visualization for
centroid size (x20), mass (x20), and substrate (x5). An exemplar sur-
face (male Gorilla gorilla) was then warped to t the estimated cong-
uration. See Methods for details.
shaped than Gorilla. It had shorter anterior, longer pos-
terior and lateral, and equal medial margins. The ante-
rior and posterior planes were at, and the medial and
lateral axes were reduced and concave. The trochlear-
medial malleolar angle was decreased (across the entire
 DETERMINANTS OF TIBIAL SHAPE 225
TABLE 8. Shape differences among taxa
Homo Pan Gorilla Pongo Hylob. Papio M. thi. M. fas. Nasalis Colobus
Whole tibia
Homo 0.0360 0.0640 0.0439 0.0502 0.0302 0.0375 0.0313 0.0373 0.0399
Pan 0 0.0336 0.0220 0.0635 0.0365 0.0266 0.0433 0.0519 0.0514
Gorilla 0 0 0.0384 0.0882 0.0626 0.0483 0.0726 0.0784 0.0789
Pongo 0 0 0 0.0615 0.0371 0.0251 0.0425 0.0509 0.0494
Hylobatidae 0 0 0 0 0.0374 0.0467 0.0314 0.0242 0.0291
Papio 0 0 0 0 0 0.0217 0.0168 0.0212 0.0229
M. thibetana 0 0 0 0 0.001 0 0.0296 0.0361 0.0358
M. fascicularis 0 0 0 0 0 0 0 0.0182 0.0156
Nasalis 0 0 0 0 0 0 0 0.002 0.0142
Colobus 0 0 0 0 0 0 0 0 0.027
Distal tibia
Homo 0.1093 0.1139 0.1525 0.1232 0.1212 0.0977 0.0992 0.1185 0.1145
Pan 0 0.0877 0.1276 0.0978 0.1186 0.0982 0.1180 0.0987 0.1284
Gorilla 0 0 0.1413 0.1146 0.1102 0.1371 0.1336 0.1163 0.1509
Pongo 0 0 0 0.1045 0.1896 0.1321 0.1510 0.1302 0.1339
Hylobatidae 0 0 0.001 0.012 0.1499 0.0951 0.1112 0.1227 0.1241
Papio 0 0 0 0 0 0.1320 0.1123 0.1162 0.1493
M. thibetana 0 0 0 0 0.035 0 0.0833 0.1237 0.1033
M. fascicularis 0 0 0 0 0.004 0 0.006 0.1127 0.1094
Nasalis 0 0 0 0 0.021 0 0.002 0 0.0874
Colobus 0 0 0 0 0.011 0 0.001 0 0.097
Medial malleolus
Homo 0.1506 0.0906 0.1948 0.1886 0.1445 0.1314 0.1568 0.1234 0.1169
Pan 0 0.1029 0.1412 0.1523 0.1697 0.1357 0.1761 0.0790 0.1347
Gorilla 0 0 0.1777 0.1782 0.1443 0.1362 0.1718 0.0931 0.0969
Pongo 0 0 0 0.1118 0.2423 0.1548 0.1682 0.1255 0.1869
Hylobatidae 0 0 0.005 0.093 0.2190 0.1421 0.1376 0.1417 0.1719
Papio 0 0 0 0 0 0.1246 0.1649 0.1546 0.1222
M. thibetana 0 0 0.001 0 0.011 0.004 0.0675 0.1179 0.1038
M. fascicularis 0 0 0 0 0.01 0 0.382 0.1614 0.1316
Nasalis 0 0.061 0.027 0.005 0.037 0 0.031 0.002 0.0991
Colobus 0 0 0.033 0 0.019 0.002 0.076 0.003 0.256
Procrustes distance between taxon means are shown above the diagonal and P values below (1,000 permutations).

Fig. 4. Visualization of distal tibial data set illustrating an exemplar surface warped to the taxon mean
landmark congurations of Papio (A), Pan (B), Gorilla (C), and Homo (D).

medial malleolar facet). The central concavity was
increased, and the trochlear-medial malleolar facet area
ratio was 4:1. The medial malleolar relative height was
reduced, with relative length equal to Pan. Finally, the
base was concave, and the central shape was at, with
the apex reduced from Pan and anteriorly displaced.
Pongo spp. demonstrated comparable robusticity and
medial angulation of the distal 1/3 of the tibia to that of
Pan, but with increased medial axial rotation, and plane
and axis of the tibiotalar joint surface comparable with
Gorilla. The trochlear shape was oval. It had an
increased medial to lateral dimension. There were
226 TURLEY ET AL.
Fig. 5. Visualization of medial malleolar data set illustrating an
exemplar surface warped to the taxon mean landmark congurations
of Papio (A), Pan (B), Gorilla (C), and Homo (D).
convex anterior, posterior and lateral margins, and a
concave medial margin. The anterior plane was
increased and posterior was at. The axis of the lateral
margin was concave and increased relative to Pan, while
the axis of the medial margin was convex but reduced.
The trochlear-medial malleolar angle was reduced from
Gorilla. The central concavity was markedly decreased,
and the trochlear-medial malleolar facet area ratio was
4:1. The relative height of the medial malleolus was
reduced, and was relatively shorter than that of Pan.
Finally, the base was at, the central shape intermediate
between that of Gorilla and Pan, with an apex reduced
from Pan but still central.
Hylobates spp demonstrated markedly reduced robus-
ticity without medial angulation of the distal tibia. It
had marked medial axial rotation. The plane of the tibio-
talar joint surface was comparable with Pan, but the
axis was comparable with Gorilla. The trochlear shape
was rectangular with an increased medial- lateral
dimension. Anterior and posterior margins were at, the
lateral convex, and the medial concave. The anterior and
posterior planes were comparable with Pan, and medial
and lateral axis comparable with Gorilla. The trochlear-
medial malleolar angle was reduced, comparable with
Pan, the central concavity was decreased, and the troch-
lear-medial malleolar facet area ratio was 3:1. The
medial malleolar relative height was reduced from Pan,
but the relative length was comparable. Finally, the base
was concave, and the central shape convex, with the
apex anteriorly displaced.
Colobus guereza demonstrated markedly reduced
robusticity without medial angulation of the distal tibia.
It had, however, marked medial axial rotation. The
plane and axis of the tibiotalar joint surface were
reduced from Hylobates. The trochlear facet shape was
oval. It had increased medial-lateral dimension at the
midpoints, with a slightly convex anterior and posterior
margin, markedly convex lateral margin, and concave
medial margin. The anterior and posterior planes were
comparable with Gorilla, as were the medial and lateral
axis. The trochlear-medial malleolar angle was reduced,
comparable with Pan, the central concavity was
decreased, and the trochlear-medial malleolar facet area
ratio was 5:1. The medial malleolar relative height was
reduced from Pan. The relative malleolar length was
greater, the base was concave, and the central shape
convex, with the apex central.
Nasalis larvatus demonstrated markedly reduced
robusticity without medial angulation of the distal tibia.
There was marked medial axial rotation, with a compa-
rable plane and axis of the tibiotalar joint surface to
Colobus. The trochlear shape was oval. There was
decreased medial lateral dimension. It also demon-
strated increased anterior and posterior convexity,
decreased lateral and medial convexity of the respective
margins compared to Colobus. The anterior plane was
at from lateral to midpoint then markedly increased
from midpoint to the anterior-medial point, and the pos-
terior plane was increased. The medial and lateral axis
was increased (medial greater than lateral) when com-
pared to Colobus. The trochlear-medial malleolar angle
was the same, while the central concavity decreased
from Colobus. The trochlear-medial malleolar facet area
ratio was reduced to 3:1, and the medial malleolar rela-
tive height was increased, comparable wiht Pan, with
the relative length was increased from Colobus. Finally,
the base was slightly convex, the central shape convex,
with the apex central.
Macaca fascicularis demonstrated reduced robusticity
without medial angulation of the distal tibia, comparable
with Colobus, but with reduced medial axial rotation
and a atter plane and increased axis of the tibiotalar
joint surface. The trochlear shape is trapezoidal compa-
rable with Homo, with a straight anterior, posterior and
lateral, and convex medial margin. The anterior plane
was slightly increased and the posterior plane slightly
convex. The lateral axis was comparable with Homo, but
the medial axis was increased. The trochlear-medial
malleolar angle was reduced from Homo, as was the cen-
tral concavity, and the trochlear-medial malleolar facet
area ratio was 3:1. The medial malleolar relative height
was comparable with Gorilla and Homo, and the relative
length comparable with Homo. Finally, the base was
at, the central shape slightly convex, with the apex
anteriorly displaced.
Macaca thibetana demonstrated moderate robusticity
without medial angulation of the distal tibia. It had
medial axial rotation comparable with Macaca fascicula-
ris. There was atter plane and increased axis of the
tibiotalar joint surface. The trochlear shape was trape-
zoidal but more rectangular, differing from Macaca fasci-
cularis, with a greater medial-lateral dimension. It had
a concave anterior, longer posterior, slightly convex lat-
eral and less concave medial margins. The anterior
plane was increased and posterior at and neutral. The
lateral axis was concave and the medial axis was convex.
The trochlear-medial malleolar angle was increased from
Macaca fascicularis but less than Homo, as was the cen-
tral concavity and the trochlear-medial malleolar facet
area ratio was 5:1. The medial malleolar relative height
was comparable with Macaca fascicularis, but the rela-
tive length less. Finally, the base was slightly convex, as
well as the central shape, with a low central apex.
Papio hamadryas anubis demonstrated decreased
robusticity, less than Macaca thibetana and comparable
with Homo, without medial angulation of the distal
tibia. It had medial axial rotation comparable with
 DETERMINANTS OF TIBIAL SHAPE
Macaca thibetana. The plane and axis on the tibiotalar
joint surface were decreased, while the trochlear shape
was trapezoidal, but more wedge shape than Homo, and
comparable with Gorilla. It had a straight anterior, pos-
terior, medial, and lateral margin. The anterior plane
was increased medially, the posterior plane at and neu-
tral, while the lateral and medial axis were concave. The
trochlear-medial malleolar angle was comparable with
Macaca thibetana. It had a decreased central concavity
and a trochlear-medial malleolar facet area ratio of 4:1.
The medial malleolar relative height was greater than
Macaca thibetana or Homo, but with a comparable
shape, while relative length was greater. Finally, the
base was slightly concave, central shape convex anteri-
orly, with an anteriorly displaced apex.
DISCUSSION
In this study, we identied the differences and similar-
ities in shape and presentation observed in the proximal
component of the talo-crural joint, the tibia, related to
substrate preference, size, and phylogeny, as well as
among the individual taxa. The implications of these dif-
ferences and similarities may provide insights into the
nature of shape change, and the selective forces acting
upon the tibia.
Differences and Similarities in Shape Related
to Biological Role
Important features differentiate arboreal and terres-
trial taxa related to their biological roles, and both pre-
sentation and shape inuence the functional
morphology. These features provide exibility and stabil-
ity on a medially placed curvilinear surface with irregu-
lar surface movement in the former, and the same
exibility and stability on a horizontally placed irregular
surface of different textures (stone, gravel, etc.) in the
latter. In arboreal taxa, there is an increase in the range
of dorsiexion and inversion, and an increase in exibil-
ity. Stability, when the foot is maximally dorsiexed, is
accomplished by increased tibial plane, while the ante-
rior portion of the medial trochlear-medial malleolar
groove and medial portion of the anterior margin were
displaced superiorly (DeSilva, 2009). Terrestrial taxa
showed lateral rather than medial axial rotation, and
the trochlear facet was trapezoid rather than oval, with
the central concavity deeper. In arboreal taxa, increasing
exibility resulted from a convex facet with a high cen-
tral apex and a convex base with antero-superior exten-
sion. The medial malleolus in terrestrial taxa provided
stability by being lower and atter, with an anterior dis-
placement of the apex, and a concave base with no supe-
rior displacement of its anterior component. Increased
robusticity closely correlated with mass, and showed
enlargement of the proximal and distal facets, with a
medial angulation of the distal tibia (differing in Mor-
phologika from curvature) among three of the Hominoid
superfamily, Pongo, Pan, and Gorilla. All three showed
moderate axial rotation, with Pongo greater, consistent
with arboreal activity and terrestrial st-walking, less
axial rotation in Gorilla accommodating its increased
mass, and less still in Pan accommodating its diverse
suite of locomotor behaviors, and diverse use of the can-
opy. Size showed shape changes due to both mass and
227
centroid size, with the effects of increased mass domi-
nated by that of Gorilla, while larger centroid size
reected those of Homo. The distal tibial facets provided
elements of stability in stance with an increase in rela-
tive medial malleolar size and a change from rectangu-
lar to apex anterior, with larger centroid size, and
widening of the trochlear-medial malleolar angle with
increased mass. Finally, with increased mass and robus-
ticity, the medial malleolar facet changed from convex to
atter with larger centroid size; demonstrated elevation
of the central base, attening of the facet face, and ante-
rior displacement of the apex. There was a more robust
tibia in hominoids than cercopithecoids, decreased AP
and increased ML dimension from cercopithecoids to
hominoids, and a slight decrease in relative surface area
of trochlear to medial malleolar facet from cercopithe-
coids to hominoids.
Differences and Similarities in Shape
Among Taxa
The inuence of size, locomotor repertoire and biologi-
cal role, likewise, inuence the shape differences and
similarities identied among the extant taxa examined.
Gorilla was an outlier in the multiple regression analy-
sis of robusticity, mass, and larger centroid size. The tra-
pezoid trochlear facet shape exceeded that observed in
more terrestrial taxa, with a wedge conguration. Tibial
medial axial rotation, and increased plane, and superior
displacement of anterior medial margin were consistent
with arboreal hominoids and colobines (Gebo, 1992).
However, Gorilla exhibits anterior displacement of a
broad atter medial malleolus suggesting accommoda-
tion to the demands of increased mass to stability in
both terrestrial activity and vertical climbing (DeSilva,
2009). This is consistent with the observation among
extant hominoids that the greatest range of motion
occurs in Pan, 55%, with a increased plane across the
entire anterior margin, followed by male Gorilla (47%)
with a atter plane laterally and increased medially,
and Homo sapiens, 48%, with a at plane across the an-
terior margin, and a signicantly greater range of
motion among female Gorilla (Latimer et al., 1987; Tur-
ley et al., 2008). Likewise, these ndings were consistent
with greater safety and stability in vertical climbing de-
spite increased body mass in Gorilla.
Pan differed from all extant hominoids in distal tibial
facets shape, with a trochlear facet plane, trochlear-
medial malleolar angle and convex medial malleolus
with tall central apex, consistent with that observed in
colobines taxa. Pan combines these hallmarks of exibil-
ity with stability in dorsiexion and inversion when ver-
tically climbing by a markedly increased anterior
margin plane and medial axis, a convex medial malleo-
lar base, reecting anterior and superior displacement of
this region (DeSilva, 2009). However, the differences
observed in metric comparisons between Pan and Homo
in relative marginal length and presumed similarities
between Gorilla and Pan were not conrmed. Tibial
articular morphology in Pan and Gorilla differed signi-
cantly (PC 1-5, and mean shape), bringing into question
the similarities proposed in their mode of locomotion
(Turley et al., 2009; Wunderlich and Jungers, 2009).
Homo, in contrast, while phylogenetically close, dem-
onstrates mean shape consistent with other terrestrial
228 TURLEY ET AL.
taxa, specically the cercopithecoids. Nevertheless,
Homo differed signicantly in shape with all taxa in all
three subsets in the pair-wise permutation test. Among
the hominoids only in Pongo and Hylobates were subsets
not signicantly different, the distal tibial facets and
medial malleolar facets. Hylobates, however, presented a
number of unique shape proles, clustering with colo-
bines and cercopithecoids (most closely with the arboreal
taxa) in PC 1-2, reecting lack of both robusticity and
tibial medial angulation, but demonstrating decreased
ML and increased AP trochlear dimensions like
Pongo, while preserving a rectangular rather than an
oval trochlear facet.
The terrestrial cercopithecoids while more robust lack
evidence of distal tibial medial angulation and demon-
strate decreased tibial medial axial rotation. Papio dif-
fered signicantly in shape with all taxa in all subsets,
with the exception of the medial malleolar facet of
Macaca thibetana, in the pair-wise permutation test.
Finally, the arboreal taxa clustered in robusticity, but
Macaca fascicularis differed in the degree of tibial
medial axial rotation, and trochlear facet and medial
malleolar shape comparable with terrestrial taxa.
Presentation, the orientation of joint surfaces in space,
affected the joint morphology observed in both homi-
noids and old world monkeys. This mainly was related
to its effect on plane. Among the hominoid taxa Hylo-
bates, lacking robusticity, showed marked medial axial
rotation, as was observed among the colobines, but with-
out distal tibial medial angulation. In the terrestrial
taxa, Papio hamadryas anubis and Macaca thibetana,
there was increased robusticity without evidence of dis-
tal tibial medial angulation, consistent with the vertical
presentation appropriate to the forces associated with
terrestrial substrate use. These results suggest a differ-
ence in tibial morphology among taxa separated since
the late Miocene (Raaum et al., 2005). Such angulation
was observed in Pongo, Pan, and Gorilla altering the
presentation of the distal articular surfaces, increasing
medial presentation to an antero-medial, arboreal sub-
strate, with less medial axial rotation. This angulation
profoundly affected plane, increasing it but requiring
only moderate tibial axial rotation. This angulation was
association with increased robusticity demonstrating the
requirement for increased size to provide the mechanical
advantage of this difference in shape. All three homi-
noids differed in the degree of medial axial rotation, but
Gorilla, the most robust, with the greatest mass, had
the most angulation. Finally, Homo, despite mild to mod-
erate robusticity like the terrestrial cercopithecoids,
lacked angulation. Lateral angulation combined with lat-
eral tibial axial rotation was observed in Homo optimiz-
ing the vertical presentation of the distal tibial articular
surface to the foot and terrestrial substrate. This combi-
nation appears to result in the square trochlear presen-
tation encountered in Homo.
Geometric morphometric analysis using GPA of land-
mark coordinates allowed examination of a number of
factors previously demonstrated to signicantly inu-
ence tibial morphology including body mass and size.
The latter complicates metric comparison of taxa, requir-
ing scaling and allometry, and restricting comparisons to
relative measurements (Latimer et al., 1987; Jungers,
1988; Ruff, 1988; Aiello and Dean, 2002; Gebo and
Schwartz, 2006; DeSilva, 2009). The former, body mass,
has been shown to correlate with articular dimensions,
complicating metric estimations and inferences concern-
ing function (Ruff, 1988; Rafferty and Ruff, 1994; Lieber-
man et al., 2001; Ruff, 2002; Pearson, 2004). Lieberman
et al. (2001), examining articular dimensions, observed
that changes in joint shape might respond to differences
in loading. They further observed that this hypothesis
required analysis of shape, not simple measurements, as
has been done in this study. The plane of the anterior
margin and axis of the medial margin of the trochlear
facet, trochlear shape, and relative length of the troch-
lear margins illustrate these points. The differences in
Pan, Gorilla, and Homo account for the stability noted
in vertical climbing in the former two despite differences
in mass and substrate preference, and terrestrial stabil-
ity in the latter two, despite differences in these same
factors.
This study examined the pure shape of the tibia in
Kendalls space (Kendall, 1984). It demonstrated both
differences and similarities among the studied taxa in
both proximal presentation and appositional articular
morphology (Conroy and Rose, 1983; Meldrum, 1991;
Stern and Susman, 1991; Lockwood and Fleagle, 1999).
The inuence of substrate preferences on variation
observed suggests that within this highly constrained
character state, homoplasty of presentation and articu-
lar shape may allow adaptation of phylogenetically dis-
tant taxa to comparable substrate demands (Lovejoy
et al., 2000; Wagner, 2001; Hall, 2005; Begun, 2007).
A critique of the methodology is reected in the fact
that the association of analogous mean shapes among
taxa with comparable substrate preferences appears
greater than the 10.9%, 6.1%, and 6.6% of Variance iden-
tied. This suggests that unlike quantitative variables,
such as robusticity, body mass, and log centroid size,
only estimates of substrate preference were available
rather than true use. Modiers such as mode of locomo-
tion (i.e., suspension, leaping, etc.), substrate availability
(specimen sites), substrate use (i.e., position in the can-
opy, trunk/terminal branches, etc.) and additional factors
may help to dene these issues, however, nothing short
of identifying the use of each individual would be compa-
rable with the quantitative variables. The small number
of specimens sampled in certain taxa, and small number
of landmarks in the medial malleolar facet subset, may
also be problematic in identifying differences among
taxa, especially in the aforementioned subsets. However,
the thin-plate splined mean shape and vector analyses
consistently identied the similarities and differences
among taxa, and their relation to the factors selected for
examination despite these issues.
Questions arise from the data observed. The tibial
medial angulation appears to be present in taxa subse-
quent to the late Miocene, with Homo, the outlier. How-
ever, the ontogeny of Homo needs to be explored. Are
the changes observed present in early infancy? Are they
due to direct genetic control of tibial shape; or are they
caused by mechanical effects of the genetic stimulus to
ambulate bipedally, vertically climb, etc., or both? What
are the ontogenetic trajectories of the entire study
group? How do these results relate to talar shape, and
how do they relate to distal presentation, the relation-
ship to the foot morphology, and substrate? Finally, how
do the shape changes identied relate to catarrhine
specimens from the fossil record?
 DETERMINANTS OF TIBIAL SHAPE
Catarrhine taxa differ in tibial morphology due to a
number of factors. In the current study substrate prefer-
ence was correlated with patterns of shape across taxa,
in spite of differences in size and taxon. Patterns of pre-
sentation differed among taxa, but were more con-
strained within superfamilies. GPA of landmark
coordinates conrmed the advantage of direct examina-
tion of shape over metric measurements as surrogates
for shape. The similarities observed in this highly con-
served joint complex among phylogenetically distant
taxa appear to reect convergent adaptations to sub-
strate, particularly of the appositional articular surfaces,
and the differences among closely related taxa may high-
light differences in substrate use and locomotor reper-
toire. These data demonstrate that proximal
presentation is closely related to size, while appositional
articular morphology is related to substrate preference.
ACKNOWLEDGMENTS
The authors are grateful to William Harcourt-Smith
and Eric Delson of the Department of Vertebrate Paleon-
tology at the American Museum of Natural History for
their assistance, Eileen Westwig of the Department of
Mammalogy at the American Museum of Natural His-
tory and Linda K. Gordon, Collection Manager at the
National Museum of Natural History for providing
access to their collection, and the Cleveland Museum of
Natural History, the P.M. Hurst Museum at the Depart-
ment of Anthropology, University of California, Berkeley,
for access to their collections, and Francis White for as-
sistance with dening substrate preference.
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