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Evidence for sexual dimorphism in the stegosaurian dinosaur Kentrosaurus

aethiopicus from the Upper Jurassic of Tanzania
Holly E. Bardenab; Susannah C. R. Maidmentc
a
Department of Animal and Plant Sciences, Alfred Denny Building, University of Sheffield, Sheffield,
United Kingdom b School of Earth, Atmospheric and Environmental Sciences, The University of
Manchester, Manchester, United Kingdom c Department of Palaeontology, The Natural History
Museum, London, United Kingdom
Online publication date: 09 May 2011
To cite this Article Barden, Holly E. and Maidment, Susannah C. R.(2011) 'Evidence for sexual dimorphism in the

stegosaurian dinosaur Kentrosaurus aethiopicus from the Upper Jurassic of Tanzania', Journal of Vertebrate
Paleontology, 31: 3, 641 651
To link to this Article: DOI: 10.1080/02724634.2011.557112
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Journal of Vertebrate Paleontology 31(3):641651, May 2011

2011 by the Society of Vertebrate Paleontology

ARTICLE

EVIDENCE FOR SEXUAL DIMORPHISM IN THE STEGOSAURIAN DINOSAUR

KENTROSAURUS AETHIOPICUS FROM THE UPPER JURASSIC OF TANZANIA

Downloaded By: [Maidment, Susannah] At: 08:31 10 May 2011

HOLLY E. BARDEN1, and SUSANNAH C. R. MAIDMENT*,2

Department of Animal and Plant Sciences, Alfred Denny Building, University of Sheffield, Western Bank, Sheffield, S10 2TN,
United Kingdom, holly.barden@postgrad.manchester.ac.uk;
2
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom,
s.maidment@nhm.ac.uk

ABSTRACTSexual dimorphism, the condition whereby males and females differ from one another physically, is one of the
most fundamental aspects of the biology of any animal. However, sexually dimorphic characters can be subtle and are mainly
related to soft tissue anatomy. They are, therefore, difficult to identify reliably in the fossil record particularly when dealing
with small sample sizes and osteology alone. The first geometric morphometric analysis of dimorphism in a thyreophoran
(armored) dinosaur shows that the femora of the stegosaurian dinosaur Kentrosaurus aethiopicus (Upper Jurassic, Tanzania)
bear a statistically significant shape difference of the proximal end, which is independent of size and is therefore proposed to
be a sexual difference. Although the disarticulated nature of the material means that intraspecific variation in other skeletal
elements, such as the enigmatic dermal armor, cannot be identified as sexual dimorphism at this time, this study provides a
methodology for further work on articulated stegosaurian specimens and has the potential to reveal additional information
regarding the palaeobiology and population dynamics of this poorly understood clade.

INTRODUCTION
Sexual dimorphism is typically defined as the physical differences between males and females of the same species (Chapman
et al., 1997). Minimally, these differences are expressed in genitalia. However, they can also be seen as differences in body size,
shape, color, and adornments such as horns, crests, or frills. Sexual dimorphism is a fundamental facet of the biology of any organism, and a far larger proportion of extant species are sexually dimorphic than monomorphic (Kaliontzopoulou et al., 2007).
Knowledge of the differences between the males and females of
a species allows insight into their behavior and evolutionary history.
Sexual dimorphism, although usually quite easily observable in living species, is very difficult to assess in extinct animals due to the lack of soft tissue preservation, often compounded by extremely small sample sizes and missing data.
Despite these problems, several studies have investigated the
phenomenon in all the major groups of dinosaur. Sexual size
dimorphisms have been suggested in the theropods Tyrannosaurus (Carpenter, 1990, Larson, 2008) and Coelophysis
(Raath, 1990), where robust and gracile morphs were considered to exist; the sauropod Barosaurus, on the basis of
growth inferred from bone histology (Sander, 2000); and the
marginocephalian Stegoceras, in which the size of the dome is
considered dimorphic (Chapman et al., 1981). Morphological differences attributed to sexual dimorphism have been identified
or suggested in the sauropods Camarasaurus and Diplodocus
based on fusion of caudal vertebrae (Rothschild and Berman,
1991), in the theropod Allosaurus based on long bone shape
(Smith, 1998), in the ornithopod Hypsilophodon where vari*Corresponding author. Present address: School of Earth, Atmospheric and Environmental Sciences, The University of Manchester,
Williamson Building, Oxford Road, Manchester, M13 9PL, United Kingdom

ation in sacral rib number was considered to be dimorphic

(Galton, 1974), within Hadrosauridae based on crest shape
(Hopson, 1975), in the marginocephalian Protoceratops based
on frill morphology (Dodson, 1976), and in the thyreophoran
Euoplocephalus, in which variation in skeletal morphology
was tentatively attributed to sexual dimorphism (Penkalski,
2001).
Many of these studies suffer from small sample sizes and, although the findings are suggestive of dimorphism, the results are
largely inconclusive. A rigorous, statistical approach to the assessment of dimorphism in Stegosauria (Thyreophora) has never
previously been carried out, although Galton (1982a, 1982b) suggested that variation in the sacral rib count of specimens of the
stegosaurs Kentrosaurus aethiopicus and Dacentrurus armatus is
related to sexual dimorphism.
The purpose of this study is to attempt to test whether dimorphism can be identified statistically through a geometric
morphometric analysis of the femora of Kentrosaurus aethiopicus. Kentrosaurus was used because the collection of material
Naturkunde
referable to this species housed in the Museum fur
(MB) in Berlin represents the largest collection of monospecific stegosaurian bones collected from a single locality anywhere
in the world (S.C.R.M., pers. observ.); the sample also represents a relatively complete ontogenetic series (the shortest femur in the collection is 31.4 cm long, whereas the longest is
71.2 cm long; Table 1), allowing effects of allometry on potential dimorphisms to be studied. Femora were chosen for the
investigation because they are in general the most abundant,
least distorted, and best preserved bones in the entire skeleton (Bonnan et al., 2008), and a large sample size (>50 individual elements) is known for Kentrosaurus. The specific questions to be tested are the following: (1) Are dimorphisms statistically identifiable in Kentrosaurus femora? (2) If so, are these
dimorphisms sexual or is variation related to ontogeny? (3) If
they are sexual, can specific morphs be assigned to females and
males?

641

642
TABLE 1.

JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 3, 2011

Specimens used in the geometric morphometric analyses arranged by size.

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Specimen number
3606
3600
3605
3604
3599
3601
3603
3561
3566
3602
3567
3562
3560
3568
3569
3558
3592
3559
3570
3590
3595
3579
3581
3578
3584
3580
3577
3583
3576
3585
3586
3591
3574
3575
3582
3594
3598
3563
3564
3565
3571
3572
3573
3587
3588
3589
3593
3596
3597

Length

Side

Shaft

Distal end

Proximal end

31.35
32.47
33.15
33.20
34.15
35.27
36.44
37.30
37.46
37.84
37.86
38.08
38.90
44.69
45.36
46.08
48.89
52.00
53.63
54.12
58.82
59.67
60.04
61.19
61.44
61.46
61.60
62.04
62.49
62.63
63.19
63.95
64.43
64.94
65.36
67.30
79.15

L
L
R
L
L
R
R
R
L
R
L
L
R
R
R
L
L
L
L
R
R
R
R
R
L
L
R
L
L
L
R
L
L
L
L
R
R
L
R
L
R
R
L
L
L
L
R?
L
L

+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ (partial)

+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+

+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+

+ (partial)
+ (partial)
+
+

+
+
+
+
+
+
+
+
+

+
+
+
+
+
+
+

Measurement a Measurement b

Group

6.81
12.07
7.79
6.90
8.09
10.06
7.64
9.28

12.44
16.30
16.72
14.11
17.70
16.25
18.37
19.49

Q
P
Q
Q
Q
P
Q
Q

7.38
8.37
5.52
8.20
11.47
10.90
8.75
7.49
10.29
16.13

17.28
17.16
17.43
14.99
17.06
16.60
17.18
17.16
18.02
22.50

Q
Q
Q
Q
P
P
Q
Q
P
P

10.39

13.24

8.74
7.94

15.82
17.56

Q
Q

+
+

For the positions of measurements a and b refer to Figure 6. Groups P and Q are different morphs: see text and Figure 7 for details.
These specimens were not included in the initial analysis because they were too incomplete.

MATERIALS AND METHODS

Naturkunde
All specimens used are from the Museum fur
and represent a sample of 49 Kentrosaurus aethiopicus femora
in varying states of completeness. There are 37 complete bones,
eight incomplete bones with both a distal and a proximal end, two
additional distal ends, one additional proximal end, and one shaft
with neither end preserved (Table 1). The material was excavated
in the early 20th century by team of German palaeontologists led
by Edwin Hennig and Werner Janensch (Maier, 2003) from the
Tendaguru beds in what was then German East Africa (DeutschOstafrika), modern day Tanzania. The bones are all from the
middle and upper saurian beds of the Tendaguru Formation
(Galton, 1982b) and are of Upper Jurassic (Upper Kimmeridgian) age (Farlow and Brett-Surman, 1999; Weishampel et al.,
2004). All reviews of the material have concluded that a single
species is represented (Hennig, 1915; Galton, 1982b; Maidment

et al., 2008). Although associated skeletons of other dinosaurian

groups were recovered from the middle and upper saurian beds,
only a single associated partial specimen of K. aethiopicus was
found, and it is now mounted, preventing it from being included
in this study (Mallinson, 2011). No other articulated or associated
specimens of K. aethiopicus were recovered and, therefore, any
dimorphism identified in femora unfortunately cannot be linked
with variation observable in other skeletal elements.
Data Collection
Photographs of the bones were taken in six different views to
investigate as much of the shape as possible. The camera was stabilized on a tripod and secured to ensure it did not move. For
each angle the bones were consistently placed to ensure that the
same view was seen each time; if necessary they were propped up
to effect a uniform orientation. Photographs of the bones were

BARDEN AND MAIDMENTSEXUAL DIMORPHISM IN KENTROSAURUS

643

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Methods

FIGURE 1. Photographs of Kentrosaurus aethiopicus femora used in

the morphometric analysis were split into sets based on their orientation.
Set 1 includes anterior and posterior views of both the complete bone
and additional photographs of the distal and proximal ends, set 2 includes
lateral and medial views of both the complete bone and additional photographs of the proximal and distal ends, and set 3 includes photographs
of end on views of the proximal and distal ends.

taken from anterior and posterior views (set 1), and lateral and
medial views (set 2; Fig. 1.); because some of the bones are incomplete, these photographs were taken both for the complete bone
and the proximal and distal ends separately in order to increase
the sample size.
Another set of photographs (set 3) were taken of the proximal and distal views of the proximal and distal ends (Fig. 1.).
Three replicates were taken of the photographs in set 1, and two
replicates of those in sets 2 and 3. Within each set replicates were
taken on different days to try to ensure that there were no errors
incurred in bone positioning or lighting. All photographs were
taken by H.E.B. to avoid potential differences in photographic
style between workers. Photographs were eliminated from sets if
the bones were too incomplete, distorted, or damaged to be of
any use in the shape analysis. Because the bones were all excavated from the same deposit, preservational biases are kept to
a minimum, although not eliminated completely. Care was also
taken to eliminate any specimens from analysis that are distorted
as a result of reconstruction during preparation.

Geometric morphometric techniques using linear measurements have a relatively long history of usage in dinosaur studies
(e.g., Chapman, 1990a, 1990b; Weishampel and Chapman, 1990;
Chinnery, 2004). Linear measurements can be very useful when
the specimens being studied grow at similar rates and in similar
directions throughout the element of interest, but this is rarely
the case (Chinnery, 2004). Measurements between homologous
points are not always able to capture all morphological changes
taking place and no matter how many linear measurements between geometrically homologous points are made, some information on shape change will be lost (Bookstein et al., 1985; Monteiro
et al., 2000; Gunz et al., 2005; Perez et al., 2006; Slice, 2007). Shape
analysis is one of a number of techniques available to quantify
overall morphological change and has proven a useful technique
to assess a number of palaeobiological problems including dimorphism (Chapman, 1990a). Shape-based methods are particularly
applicable when smaller amounts of material are available than
would be required for a more traditional geometric method based
on linear measurements (Chapman, 1990a) and they allow subtle
shape changes to be visualized (Perez et al., 2006); we therefore
use landmark-based shape analysis in this study.
The landmarks used to analyze the shape of the femora were
all type 2 (Bookstein, 1991), i.e., points defined by local properties such as maxima of curvature. Landmarks were selected to
be homologous only to the extent that they could be consistently
found within the sample studied, because no phylogenetic analysis was attempted in this investigation. The landmarks that were
chosen consisted of all the points on the samples that could be reliably and consistently located on each bone, although the shaft
midpoints were also included and were found on the placement
of a comb (see below). The bones have relatively few distinct
anatomical points on which to place landmarks that can be consistently identified on all specimens, so semilandmarks were also
used to represent the shape of the bones outline or profile (Fig.
2, Table 2).
Although landmarks are geometrically homologous points
(Gunz et al., 2005), from a biological standpoint, curves are
homologous between individuals as well, even if points along
them are not (Perez et al., 2006). Semilandmarks are points
placed along a biologically homologous curve spaced to minimize their bending energy, which reduces the potential for overinterpretation of the deformation grid. Once the bending energy
has been minimized, the semilandmarks can be treated as homologous points without artefact (Gunz et al., 2005). Semilandmarks permit the representation of the overall shape of a biological structure and incorporate information lost when landmark or
linear data alone are used (Perez et al., 2006).
In order to facilitate the placement of landmarks and semilandmarks, both fans and combs were placed on photographs of the
bones using the IMP software MakeFan6 (Sheets, 2002; Fig. 2.).
Fans and combs are useful tools for providing guidance in the
placement of semilandmarks on areas of indistinct shape. When
anchored between specified landmarks, fans give points in a circle or semicircle at regular angles, whereas combs produce linear
points at regular distances. Because the points given by fans are
not regularly spaced, they are not suitable for eigenshape methods and therefore can only be used to generate semilandmarks
(Sheets et al., 2006). They are commonly used in studies involving bones or other samples that make use of semilandmarks over

curves (Sheets et al., 2004, 2006; Martinon-Torres

et al., 2006).
Where combs were used, they were positioned between the most
proximal point of the femoral head and the approximate midpoint between the lateral and medial condyles. Fans were also
consistently placed: for example, on the proximal end of the anterior view, they were placed between the most proximolateral
point of the greater trochanter and the most medial point of the

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644

JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 3, 2011

FIGURE 2. A, anterior view of the complete femur of Kentrosaurus aethiopicus (MB R.3574) showing the placement of landmarks (closed circles)
and semilandmarks (open circles) used in the geometric morphometric analysis of its shape; B, lateral view of the complete femur (MB R.3561)
showing the placement of landmarks and semilandmarks. See details of the landmark placement on both views in Table 2.

femoral head. All landmarks were digitized using the software

tpsDig 2.12 (Rohlf, 2008). Relatively fewer semilandmarks were
used in the original analyses; however, preliminary investigations
suggested that significant shape variation existed in the proximal
end of the bone in the anterior view, and so more were employed
in order to fully represent the shape of the bone.
Within each set of photographs, either those of the left or
right femora were horizontally inverted according to whichever
was in the minority within the set. The semilandmarks were
then allowed to slide in order to minimize their bending energy (Bookstein, 1991) using the software tpsRelw 1.46 (Rohlf,
2008). Further analyses were carried out using the software MorphoJ 1.00 (Klingenberg, 2009), the first of which was a generalized Procrustes analysis. This scales, rotates, and translates the
specimens to remove the effects of size and orientation (Slice,
2007). Deformation grids, principal components analyses (PCAs)
and canonical variates analyses (CVAs) were used to visualize
any differences in shape and multivariate analysis of variance
(MANOVAs) were used to determine differences between the
means of any groups found within the sample. t-Tests were used
to identify specific shape differences, and a regression was used to
investigate the effects of allometry on the shape differences that
were discovered.
Clusters were initially identified visually from the PCA plots,
after which a k-means cluster analysis was used to discover
whether these clusters were valid. k-means clustering separates
data into a number of groups chosen by the investigator to represent, for example, separate populations, geographical areas, or

TABLE 2. Definitions of the landmarks used to analyze the shape of

the anterior and lateral view of the complete Kentrosaurus aethiopicus
femora, shown in Figure 2.
Complete boneanterior view
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.

Most proximal point of the femoral head

Most medial point of the femoral head
Midpoint of the medial shaft
Most medial point of the medial condyle
Most distal point of the medial condyle
Midpoint between the medial and lateral condyles
Most distal point of the lateral condyle
Most lateral point of the lateral condyle
Midpoint of the lateral shaft
Most lateral point of the lateral rugosity
Most proximolateral point of the greater trochanter

Complete bonelateral view

1. Most proximal point of the femoral head
2. Anterior inflection point between the femoral head and the
anterior trochanter
3. Most anterior point of the anterior trochanter
4. Midpoint of the anterior shaft
5. Most anterior point of the medial condyle
6. Point of inflection between the medial and lateral condyles
7. Most distal point of the lateral condyle
8. Most posterior point of the lateral condyle
9. Midpoint of the posterior shaft
10. Most posterior point of the greater trochanter
11. Posterior point of inflection between the greater trochanter and
the femoral head

BARDEN AND MAIDMENTSEXUAL DIMORPHISM IN KENTROSAURUS

sexes. The analysis attempts to group data points into this given
number of groups so that intra-group variation is greater than
inter-group variation (MacQueen, 1967). In this study k-means
analyses were performed in PAST (v2.01; Hammer, 2010), in
which the data were separated into two k-groups.
RESULTS
No significant clustering was seen on the PCA plots of the complete bone (Fig. 3) or distal ends in sets 1 and 2, and none in the
distal or proximal ends in set 3. However, clustering was seen on
the PCA plots of both the lateral (set 2; Fig. 4) and anterior (set
1; Fig. 5) views of the proximal end.

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Proximal End in Lateral View

A MANOVA was performed on the groups identified from the
clustering observed in the PCA plot (Fig. 4). This showed that
they differed significantly in shape (F = 12.98, P 0.0001) but not
in (centroid) size (F = 0.37, P = 0.55). A difference in bone shape
but not in size indicates that the dimorphism is not ontogenetic
and is probably sexual; however, the deformation grid shows that
the shape differences are widespread over the proximal end and
therefore hard to define, making them diagnostically ineffectual
for separating the sexes.
Proximal End in Anterior View
A MANOVA performed on the groups identified by the PCA
plot (Fig. 5) showed that they differed significantly in shape (F
= 12.59, P 0.0001) and also in (centroid) size (F = 9.00, P =
0.0071), indicating that the groups are most likely to represent
ontogenetic rather than sexual dimorphs. The sample was then
split into juveniles and adults using the distance between the most
proximolateral point of the greater trochanter and the most medial point of the femoral head (shown as measurement b on Fig.
6A) as a measure of size (the length of the complete bone could
not be used as some of the proximal ends in the sample were from
incomplete specimens, and a linear regression indicated that this
measurement was an effective measure of the size of the complete bone [F = 20.87, P = 0.0008]).
Two groups were identified in a PCA plot of the adult specimens in the sample (Fig. 5, Table 1) that differed significantly
in shape (MANOVA, F = 9.19, P 0.0001) but not significantly
in (centroid) size (F = 0.03, P = 0.87). The two groups determined by k-means cluster analysis separated the data into exactly
the same groups as were chosen on visual inspection of the PCA
plot (Table 3). The separation of the two adult groups and the
juveniles is more easily seen on the canonical variates analysis
(Fig. 5B.). The deformation grid of the first principal component
(PC) (Fig. 6B) showed that the major difference in shape between the two adult groups occurred in the distance between the
most lateral point of the lateral rugosity and the most proximolateral point of the greater trochanter (between landmarks 1 and
2, shown as measurement a on Fig. 6A). This shape difference
accounted for 57.9% of the total variation. A two-tailed, twosample t-test on this distance between the two groups showed a
significant difference (t = 5.11, P = 0.0002); the same test was performed on measurement b (Fig. 6A), showing that there was an
insignificant difference in the size of the bones of the two groups
(t = 0.76, P = 0.46). A linear regression was carried out on measurements a and b (Fig. 6A), which indicated that they are not
significantly allometrically related (F = 4.91, P = 0.044).
Because a statistical difference between the two groups had
been established, other specimens were included in the investigation that had previously been excluded from morphometric analysis because they were too incomplete. Measurements a and
b were taken from specimens MB R 3575, 3576, 3580, 3583,
and 3591 using tpsDig 2.12 (Rohlf, 2008; Table 1). Another two-

645

tailed, two-sample t-test was carried out that included the measurements from the new specimens. This also showed a significant
difference in measurement a (t = 5.81, P = 0.0001) between the
two groups but not in measurement b (t = 0.466, P = 0.647).
There is a 49.33% difference in measurement a between femora
in groups P (those with a larger measurement a) and Q (those
with a smaller measurement a) (Table 1).
DISCUSSION
A large amount of individual variation is observed in the shape
of the complete femora of Kentrosaurus. However, not all of this
variation is definable as a distinct series of dimorphisms. In contrast, a distinct shape dimorphism is identifiable at the proximal
end of the femur in anterior view. This is not coupled with a significant difference in femoral size and, therefore, is unlikely to be
related to allometry or ontogeny. Instead, we propose that it is
most likely a sexual dimorphism. Further evidence for this is provided by the fact that the dimorphism is only detectable in large
femora, suggesting that it develops at sexual maturity.
The main difference in shape is the distance between the most
lateral point of the lateral rugosity and the most proximolateral
point of the greater trochanter (measurement a, which effectively corresponds to the size of the greater trochanter; Figs. 6A,
7). The main shape changes that occur on PC1 of the anterior
view of the total bone show a general trend from a more gracile
form to a more robust one with a more pronounced and better developed greater trochanter, a more distinct femoral head
and larger distal condyles; however, these data include the complete sample whereas the plot of the proximal end only includes
the adults. The difference shown in the PCA of the total bone
is therefore more likely to show ontogenetic changes associated
with general bone growth; the same is true for PC1 on the posterior view of the total bone.
The dimorphism indentified in the sample is not only seen on
the PCA plot but is also confirmed with both a k-means cluster
analysis and a t-test. k-Means cluster analysis has been used to
great effect in other geometric morphometric studies of sexual
dimorphism in living animal populations (Valenzuela et al., 2004;
Gonzalez et al., 2009). Such studies do, however, have the advantage of being able to re-test the shape variation against actual
sex within the sample in order to validate the dimorphism. In
such studies, the groups determined by k-means analysis haven
been shown to be up to 97% accurate in assigning sex (Valenzuela et al., 2004), although this does vary depending on the type
of animal studied and the skeletal element used. k-Means clustering requires the investigator to determine the number of groups
into which the data are assigned. Here, we make the assumption
that there are two groups. This is an unavoidable area of potential bias in our analysis, but visual inspection of the data suggested
that the assignment of two groups was most parsimonious for our
data.
Being a least squares method, generalized Procrustes analysis is at risk from the Pinocchio effect (Chapman, 1990c): a
distortion of the data as a consequence of a few landmarks being significant outliers. The analysis minimizes the sum of the
squared distances and spreads the variation of the outliers to
other landmarks, smearing it across the whole shape (Zelditch
et al., 2004). To overcome this problem, some authors (Chapman,
1990b; 1990c) have suggested the use of resistant-fit methods that
are less sensitive to outlying landmarks and provide more accurate representations of the shape variation. Resistant-fit methods, however, do not produce Procrustes distances and because
their residuals are not homologous, they cannot be compared between samples, thus precluding the use of these data in subsequent statistical analyses (Bookstein, 1996). It is possible that our
landmarks 1 and 2 produce a Pinocchio effect; however, this
would cause the variation between them to be underestimated

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FIGURE 3. Plots showing the first against second principal components for the data from sets 1 and 2. A, total anterior view (set 1); B, total posterior
view (set 1); C, total lateral view (set 2); D, total medial view (set 2). The shape of the bones at the extreme of each axis is shown as a wireframe.

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BARDEN AND MAIDMENTSEXUAL DIMORPHISM IN KENTROSAURUS

647

FIGURE 4. A plot of the first against the second principal components

for the morphometric analysis on the lateral view of the proximal end of
the Kentrosaurus femora. Each point represents one specimen; the separation of the individuals into statistically distinct groups is shown by
means of the circle on the graph. Wireframes are present to show the
shape of the bone at the extremes of the axes.

and any variation seen would therefore be under-representative

of the true level of variation within the sample. The shape variation observed in our results is therefore likely to be real and not
due to the effects of a few outlying points.
It could be argued that the dimorphism indicates a specific difference, but there is currently no other evidence to suggest that
more than one species of Kentrosaurus is present in the fauna
(Hennig, 1915; Galton, 1982b; Maidment et al., 2008). The fact
that the dimorphism can only be identified in larger femora also
tends to suggest that this is not the case. It is possible that, once
sexual maturity had been attained and growth slowed, an increase in robusticity of limb elements occurred with increasing
age. However, because the sample divides into two distinct clusters not correlated with size (Fig. 5), we consider that sexual dimorphism is more likely, although an independent test of age,
such as a histological study and identification of external fundamental systems, would help to rule out this possibility.

FIGURE 5. A, a plot of the first against the second principal components for the morphometric analysis on the anterior view of the proximal
end of the Kentrosaurus femora of the adult specimens in the sample.
Each point represents one specimen; the separation of the individuals
into groups is shown by means of the circle on the graph. Those individuals within the circle are in group P and those outside it are in group
Q. Wireframes are present to show the shape of the bone at the extremes
of the axes. B, graph showing the first against the second canonical variates from the same analysis of the whole sample. The three groups separated on the graph represent the adults, split into groups P and Q, and
the juveniles.

Identifying Sex
Although establishing whether a species is sexually dimorphic
is important, biologically it is far more useful to know which
morph is male and which is female. In paleontological investigations this is usually where fact merges with speculation, because
it is very rare to have enough information to make an informed
decision on gender. Previous studies have found similar dimorphisms in femoral structure. Raath (1990) found a dimorphism
in the dimensions of the greater trochanter in the femora of the
theropod Coelophysis rhodesiensis; the two groups represented

gracile and robust morphs, of which the robust form was cautiously proposed to be female. A bimodal distribution was identified in the femoral structure of the semi-aquatic reptile Champsosaurus, thought to be potentially due to different locomotory
regimes imposed on males and females associated with females
leaving the water to lay eggs (Katsura, 2004). A femoral sexual
dimorphism was also found in Tyrannsoaurus rex (Larson, 2008),

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TABLE 3. k-Means cluster analysis of the anterior view of the proximal
femoral head.
Specimen number

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3570
3572
3574
3578
3579
3581
3582
3584
3585
3586
3587
3588
3590
3594
3595
3598

FIGURE 6. A, anterior view of the proximal end of a Kentrosaurus

aethiopicus femur (MB R.3574) showing the placement of landmarks
(closed circles) and semilandmarks (open circles) used in the geometric
morphometric analysis of its shape. 1, most lateral point of the lateral
rugosity; 2, most proximolateral point of the greater trochanter; 3, most
proximal point of the femoral head; 4, most medial point of the femoral
head. Measurement a (the distance between landmarks 1 and 2) represents a probable sexual dimorphism; measurement b (the distance between landmarks 2 and 4) is an effective measure of the total size of the
bone. Measurements a and b for each specimen within the sample are
shown in Table 1. B, a deformation grid of the first principal component,
showing the main changes in the shape of the proximal end of the femur
in anterior view; landmarks used are the same as described in A (above).
The main change that can be seen occurs in the distance between landmarks 1 and 2, represented by measurement a.

k-Means clustering

Group

2
1
1
1
2
2
2
2
2
2
2
2
1
1
2
1

Q
P
P
P
Q
Q
Q
Q
Q
Q
Q
Q
P
P
Q
P

where morphs were again classified as robust and gracile. In this

case, the robust form was identified as female based on other
skeletal evidence, including pelvic dimensions and the presence
of fused caudal vertebrae. Further support for this assessment
was provided by the discovery of medullary bone in T. rex femora
(Schweitzer et al., 2005) that clustered significantly with the robust forms from the previous investigation. Medullary bone is
produced by female birds at certain times during the reproductive cycle; it acts as a calcium reservoir to aid in the formation of
egg shells (Dacke et al., 1993). Currently the only way to reliably
assign gender to sexual morphs in the fossil record is by identification of medullary bone or by the discovery of a fossil that was
giving birth or pregnant at the time of death.
Another potential means of inferring sexual identity in dinosaurs is to examine the patterns of sexual dimorphism in their
extant phylogenetic bracket. Patterns of sexual dimorphism are
very varied in birds. Some species are monomorphic, such as the
European swift (Apus apus; Owens and Hartley, 1998). In some
taxa, such as raptors (Raath, 1990), females are larger than males,
whereas in others, such as the musk duck (Biziura lobata; McCracken et al., 2000), males are far larger than females. Even
among basal birds, such as ratites and tinamous, sexual size dimorphism is variable. Females are larger than males in kiwis
(Apteryx), southern cassowaries (Casuarius casuarius), the extinct moa Dinornis (Bunce et al., 2003), emus (Dromaius; Malony and Dawson, 1993), and tinamous (Tubano and Bertelli,
2003), whereas in ostriches (Struthio) males are larger than females (Handford and Mares, 1985). In crocodilians, males tend to
grow faster than females and to a larger size, but show no other
obvious external differences (Grigg and Gans, 1993). Patterns of
sexual size dimorphism are extremely varied in the extant phylogenetic bracket of dinosaurs and so are of limited use for determining sexual identity.
In this study, the specific shape difference between the two
morphs concerns the dimensions of the greater trochanter, a site
of attachment of the puboischiofemoralis externus ( = obturator)
musculature in crocodiles and birds (Romer, 1927; Hutchinson
and Gatesy, 2000). Because the relative size of a muscle attachment point can indicate the size of the attaching muscle (Gatesy,
1990), it is probable that one morph had a greater muscle mass
than the other. This difference in muscle mass between the sexes
could be explained in a number of ways: (1) A greater muscle
mass would have allowed the support of a greater body mass, indicating that one sex was larger than the other. (2) Because the
puboischiofemoralis externus is a femoral protractor and therefore related to locomotion in crocodiles (Gatesy, 1997), differences in muscle mass could indicate that the different sexes had
different locomotory regimes, for example. (3) A greater muscle

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649

FIGURE 7. Examples of the proximal end morphology of Kentrosaurus aethiopicus from groups P and Q. A, example of a femur from group P
(MB R.3578) showing a relatively larger distance between the most lateral point of the lateral rugosity and most proximolateral point of the greater
trochanter than those in group Q; B, example of a femur from group Q (MB R.3585). Scale bars equal 10 cm.

mass could be an adaptation related to the mechanics of copulation: the male could have required a greater muscle mass in order
to mount the female, or conversely the female may have required
the greater muscle mass in order to give extra support while being mounted. We conclude that there is not enough evidence to
attempt to identify either group as male or female.
Sex Ratio
The dimorphism identified in Kentrosaurus separated the sample in a ratio of approximately 2:1 in the original analysis and
exactly 2:1 with the inclusion of more specimens. By far the
most common form of polygamous mating system in vertebrates
is polygyny, whereby males mate with more than one female
(Marlowe, 2003), sometimes defending harems from other males.
This pattern is seen in the American crocodile (Crocodylus acutus), American alligator (Alligator mississippiensis), and Nile
crocodile (Crocodylus niloticus) (Garrick and Lang, 1977); each
male mates with more than one female and usually defends a
breeding territory. Patterns in the mating systems and sex ratios
of birds are far more variable. In this study, femora with morphology belonging to group Q are twice as common as those belonging to group P, which might suggest that group Q represent
females. However, caution must be taken in interpreting the specimens in the context of a population because they are probably
not the result of a mass death assemblage and so may not adequately represent population structure. The animals are instead
thought to have gathered in the area gradually due to a drought,
and died there as they eventually overpopulated the area and exhausted its resources (Aberhan et al., 2002).
CONCLUSION
Kentrosaurus aethiopicus was dimorphic in the dimensions of
the proximal end of its femora. This dimorphism is independent of size and, therefore, most likely represents a sexual dimorphism. The nature of the dimorphism means that one morph
was more robust, i.e., supported a greater muscle mass, than the
other. Other studies that have investigated sexual dimorphism
using femora in dinosaurs have also identified shape differences

in the proximal femoral head, although all have been based on

measurement data or qualitative observation rather than shape
analysis (e.g., Weishampel and Chapman, 1990; Smith, 1998, Larson, 2008). Raath (1990) found similar differences, albeit qualitatively, in greater trochanter size in the theropod Coelophysis as
we report here. It is possible that sexual size dimorphism in hip
musculature is a feature common to all dinosaurs; this generalized observation warrants further investigation. Identification of
sexual dimorphism in extinct animals is crucial to our understanding of population structure and dynamics, diversity, and evolutionary history and this finding sheds light on the paleobiology
of this poorly known clade of ornithischian dinosaur. This is the
first study to provide statistical evidence for sexual dimorphism in
stegosaurian dinosaurs; further research could focus on taxa for
which there are well-preserved, articulated specimens, for example, the North American genus Stegosaurus. A similar shape dimorphism, if identified, could then be linked to variation throughout the skeleton; in particular, shape and size variations in the
enigmatic dermal armor possessed by stegosaurs.

ACKNOWLEDGMENTS
This work forms part of the MBiolSci thesis of H.E.B., which
was supervised by C. Wellman (University of Sheffield) and
S.C.R.M., and for which the Department of Animal and Plant
Science, University of Sheffield, provided logistical support and
funding for overseas travel. Thanks to D. Schwarz-Wings (Mu Naturkunde) for access to specimens in her care. N.
seum fur
Gidaszewski (University of Sheffield) and M. E. H. Jones (University College London) provided help and advice regarding geometric morphometric techniques and figures. We thank delegates
at the Symposium on Stegosauria, held at the Sauriermuseum,
Aathal, Switzerland, in June 2009, for constructive discussion and
comments. P. M. Barrett (Natural History Museum, London), K.
Padian and S. Werning (University of California, Berkeley), M.
E. H. Jones (University College London), reviewers P. Galton
(University of Bridgeport) and R. Chapman, and handling editor
R. B. Holmes (University of Alberta) provided extremely helpful
comments on earlier drafts of the manuscript.

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