10. Sauropod Tracks – a geometric morphometric
study
Luis Azevedo Rodrigues, Vanda Faria dos Santos
Museu Nacional de História Natural (MNHN), Universidade de Lisboa, Rua da
Escola Politécnica, 58, P-1250-102 Lisboa, PORTUGAL, e-mail:
lmrodrigues@fc.ul.pt
10.1 ABSTRACT
Geometric morphometrics are used to characterize shape variations in different Sauropo-
domorpha ichnotaxa and unclassified ichnites. Ten landmarks were collected from each of
30 specimens. Landmark configurations were superimposed, and residuals were modeled
with the thin-plate spline interpolating function (to visualize shape changes). This group of
techniques allows to discriminate tendencies in shape changes (providing quantitative de-
scriptors).
The multivariate analysis of shape variables on the centroid size indicates the absence of al-
lometry in our sample of Sauropodomorpha tracks.
Keywords: Sauropoda pes tracks; Brontopodus; Geometric morphometrics; allometry;
Relative Warps
10.2 INTRODUCTION
Application of Geometric Morphometric (GM) techniques in the ichnological
record haven’t received much attention. Some applications have been made on di-
nosaur tracks particularly the Theropoda and Ornithopoda ichnological record
(Rasskin 1995; Rasskin et al. 1997). This work presents the first GM study on the
Sauropodomorpha ichnological record. Here we constrast a descriptive study
based on four sauropod ichnological morphotypes with a geometric morphometric
approach.
The discovery and documentation of many new sauropod tracksites in Portugal
over the last ten years have yielded valuable information to better understand
sauropod manus and pes prints morphologies. Nevertheless, well-preserved sauro-
pod manus and pes prints are still rare in the general fossil record.
Until 1990 a small number of well-preserved specimens were known world-
wide and few ichnogenus were considered valid scientific names. Middle Jurassic
sauropod ichnites from Morocco were described as Breviparopus taghbaloutensis
(Dutuit and Ouazzou 1980).

130 Luis Azevedo Rodrigues, Vanda Faria dos Santos
Brontopodus birdi was named on the basis of well-preserved sauropod track-
ways from Albian carbonates from Texas (Farlow et al. 1989). Parabrontopodus
mcintoshi was proposed from narrow-gauge Upper Jurassic sauropod trackways
(Lockley et al. 1994a)
The record of the ichnogenus Brontopodus is worldwide distributed - Portugal
(Lockley et al. 1994a,b), Croatia (Mezga and Bajraktarevic 1999), Switzerland
(Meyer 1993), Spain (e.g. Moratalla 1993), Poland (Gierlinski 2002), United
Kingdom (Romano et al. 1999), China (Lockley et al. 2002), South Korea (e.g.
Lim et al. 1994), Australia (e.g. Thulborn et al. 1994), USA (Farlow et al. 1989).
More recently, Lower Cretaceous sauropod footprints from Croatia were de-
scribed and named as Titanosaurimanus nana (Dalla Vecchia and Tarlao 2000).
In the present study were used Sauropodomorpha footprints outlines from sev-
eral works (Table 1).
Most of the sauropod footprints are oval- or egg-shaped without diagnostic
digit impressions, due to inadequate substrate conditions, and further unfavorable
conservation factors (e.g. Leonardi 1987; Lockley 1991; Gatesy et al. 1999; Gar-
cia-Ramos et al. 2000; Nadon 2001). Nevertheless, several of these poorly pre-
served ichnites have received formal names despite the fact that other well-
preserved specimens have not (Lockley et al. 1986).
Up to now, no Geometric Morphometric analysis of shape variation in a sam-
ple of Dinosauria – Sauropodomorpha footprints of the world ichnological record
has been conducted and just only the preliminary results of the application of this
methodological approach on 22 specimens were presented (Rodrigues and Santos
2003). The aim of this paper is to discuss the contribution of the Geometric Mor-
phometrics analysis to improving the discrimination of Sauropodomorpha foot-
prints and, possibly, to improving the characterization of ichnological shape varia-
tion.
In this paper we use GM analysis on pes prints attributed to Sauropoda and
other marks attributed to Prosauropoda footprints in order to provide a contribu-
tion to Sauropodomorpha footprints discrimination. We chose a total of 30 Sauro-
podomorpha pes tracks from the world ichnological record (range of standard
length 5.8–94.0 cm) (see Table 1).

10.3 Materials and methods
10.3.1 Samples
Good general preservation and presence of, at least, four digit impressions de-
termined selection of specimens. With these selection criteria, we tried to reduce
the taphonomical bias.

10. Sauropod Tracks – a geometric morphometric study 131
The descriptions of sauropod tracks were based on several features characteris-
tics of Sauropoda autopodia. Concerning pes prints the following distinctive fea-
tures were characteristic:
 subcircular/suboval/subtriangular shape of the pes print with asymmetrical ex-
panded proximal portion (entaxonic);
 outward rotated;
 four or five digit impressions usually outward rotated or laterally oriented;
 strongly curved and usually triangular impressions;
 claws on digits I, II, III.
We grouped the specimens into four morphotypes, based on the ichnotaxonomy
proposed by different authors in the literature. Morphotype 1 (MT1) – this mor-
photype gathers Brontopodus birdi/Brontopodus sp.; Morphotype 2 (MT2) –
Brontopodus aff. B. birdi / Brontopodus type / Brontopodus; Morphotype 3 (MT3)–
Prosauropoda ichnites (Tetrasauropus/ Pseudotetrasauropus/ Paratetrasauro-
pus); Morphotype 4 (MT4) – miscellaneous and unidentified ichnites.

Fig. 1. Examples of specimens used in this study – Specimen 1 (Morphotype 1), Specimen
2 (Morphotype 2), Specimen 8 (Morphotype 4), Specimen 12 (Morphotype 1), Specimen
26 (Morphotype 2) and Specimen 18 (Morphotype 3). Parataxonomy and references in Ta-
ble 1. IV – fourth digit. (Adapted from Lockley et al. 1994a, Santos et al. 1994, Thulborn
1990).
10.3.2 Obtaining Landmarks coordinates
Due the inherent characteristics of the materials analyzed in this study, the type of
landmarks applied were Type III (Bookstein 1991).
Silhouettes and photos of Sauropodomorha footprints in literature were used
and digitized with Hp 5470C scanner.
Images were treated digitally (digital clearness) using Paint Shop Pro 7.0 (Jasc
Software 2002).
We assumed that all specimens were left pes. When only right pes existed, the
specimens were reflected (mirror effect) using Paint Shop Pro 7.0 (Jasc Software
2002).
The coordinates of the specimens were determined with TpsDig 1.37 (Rohlf
2003a). Since we used figures from different literature sources, they presented dif-
ferent sizes. In order to correct this we used scale factor in every specimen.

132 Luis Azevedo Rodrigues, Vanda Faria dos Santos
Some of the specimens studied have been measured in terms of length and
width, applying the measure tool on TpsDig 1.37 (Rohlf 2003a). This procedure
was applied because those measurements are not mentioned in the literature.
10.3.3 Description of landmarks
1- maximum hypex of digit I; 2 - hypex between digits I and II; 3 - maximum hy-
pex of digit II; 4 - hypex between digits II and III; 5 - maximum hypex of digit III;
6 - hypex between digits III and IV; 7 - maximum hypex of digit IV; 8 – intersec-
tion point between a perpendicular line (from mid point of landmark 7 and 9) and
the ichnite contour; 9 – most posterior point of the print considering its axis; 10 -
intersection point between a perpendicular line (from mid point of landmark 1 and
9) and the ichnite contour .
All ichnological terms and definitions follows Leonardi (1987) and Thulborn
(1990). The long axis of the footprint employed in the landmarks 8, 9 and 10 fol-
lows the definition of Leonardi (1987).


Fig. 2. – Orientation terminology and position of the 10 landmarks used in this study.
Landmark 1 and 9 are in the anterior and posterior region of the track, respectively; land-
mark 8 and 10 are in the lateral and medial region of the track, respectively
The mid points between landmarks 1 and 9 and between 7 and 9 were calcu-
lated. Marking of a perpendicular line to the referred mid points and the point of
intersection of that line with the contour of the footprint. These calculations were
performed with Microsoft Visio 2000 (Microsoft Corporation 2000).
A full, detailed, mathematical description of the GM methodology used in this
study is outside the range of this paper. Theoretical background of these meth-
dologies are reviewed in different literature sources (e.g. Bookstein 1989a, b,
1990, 1991; Rohlf and Marcus 1993; Marcus et al. 1996; Rohlf and Bookstein
2003).

10. Sauropod Tracks – a geometric morphometric study 133
For each specimen, centroid size and weight matrix (with both uniform compo-
nents appended) were computed. The weight matrix is the matrix of the partial
warp scores. Centroid size was tested for differences by single classification
analysis of variance (ANOVA) (Sokal and Rohlf 1995). All specimens were
scaled to unit centroid size before alignment by the method of Generalized Pro-
crustes Analysis (GPA) superimposition.
10.4 Relative warp analysis
The coordinates of all aligned specimens were used for thin-plate splines relative
warp analysis (Bookstein 1991; Rohlf 1993).
The Relative Warps (RW) analysis was performed with the scaling option α=0
(Rohlf 1993) that weights all landmarks equally, with the uniform component in-
cluded - complement method (Rohlf and Bookstein 2003).
Relative warps analysis corresponds to a Principal Components Analysis of the
covariance matrix of the partial warp scores, which are different scales of a thin-
plate spline transformation of landmarks. The thin-plate spline is a smooth inter-
polation function that computes and visualizes transformations of Cartesian Coor-
dinates in a way similar to D’Arcy Thompson’s transformation grids (Thompson
1917). A rectangular grid is projected over Procrustes aligned landmark configura-
tions and the bending of the grid visually depicts the difference in landmark loca-
tions between two configurations
The columns of the weight matrix represent the shape variables (Partial warps),
being the last two columns the uniform shape components (Unif X, shearing, and
Unif Y, stretching along the major axis of the consensus configuration). The first
n-2 columns characterize more localized shape components (non uniform shape
components).
10.5 Multiple Regression analysis
Centroid size, the square root of the sum of the squared distances between all ho-
mologous landmarks and the center of gravity of the landmarks, is commonly used
as general size measure in geometric morphometrics.
To explore the existence of size allometry (i.e. shape change as a function of
size), a multivariate regression of the weight matrix (with uniform components
appended) onto log centroid size was performed. The log of centroid size was used
as our size variable because most of shape change occurs early in ontogeny (e.g.,
Zeldich et al. 2000).

134
10.6 Software
Procrustes superimposition, weight matrix, graphical material and centroid size
were performed using TPSRelw 1.35 (Rohlf 2003b); TPSRegr 1.26 (Rohlf 2003c).
Statistical analysis and scatterplots – SPSS 10.0 (SPSS Inc., 2000).
10.7 RESULTS
Centroid size was tested for differences among specimens by ANOVA (F= 4.126,
p< 0.05) and was significant. The centroid size is moderately correlated with the
uniform component x (r=-0.379, p<0.05) and not correlated with uniform compo-
nent y.
10.7.1 Relative warps analysis

Fig. 3. Scatterplot of mean consensus configuration with individual specimens su-
perimposed by Generalized Procrustes Analysis (GPA)

There is large variability in most of the landmarks as visualized in Fig. 3. In land-
marks 8, 9 and 10 the observed variability is mostly latero-medial.
The first three relative warps account for 66.46% of the total variation of the
specimens. RW1 accounts for 35.52% of total shape variability. There is a signifi-
cant correlation between centroid size and the first relative warp (r=0.404, P
<0.05).
RW2 accounts for 18.40% for shape variability and is not correlated with cen-
troid size. RW3 explains 12,54% of shape variability and is not correlated with
centroid size.
The distribution of the four morphotypes is presented in a scatterplot of Rela-
tive Warp 1 (RW1) and Relative Warp 2 (RW2) (Fig. 4).

10. Sauropod Tracks – a geometric morphometric study 135
Distribution of specimens presents some tendencies: specimens 16, 17 and 18
(Prosauropoda origin) clearly separated from other specimens; specimens 1, 2,
12, 24 and 26 are morphological related despite their different morphotype classi-
fication; Croatia specimens (27, 28, 29 and 30) are closely associated with the ex-
ception of specimen 30. This could be explained as probable misidentification of
digit polarity. All other specimens present a distribution very similar with the con-
sensus and without clear grouping.
Patterns of shape change along the two first relative warps are illustrated in
Fig.5. Most of the variance from the consensus, along RW1 axis (negative to posi-
tive deviations), is due to the rotation of digits from an inward (medial) position to
an outward (lateral) position. Similar tendency in relative shape change is ob-
served in the heel region. This shape variation along this axis is also a conse-
quence of a medial bending. In addition, we detected an antero-posterior shorten-
ing associated with digit rotation (both outward and inward). The correlation
between centroid size and the first relative warp support this size shift.
RW2 differences from the consensus (negative to positive deviations) are due:
1- reduction in relative length of digits I and II; 2 – digits I, II and III becoming
narrow.

Fig. 4. Scatterplot of RW1 and RW2 of the specimens and respective Morphotypes

136


Fig. 5. Shape changes depicted by the RW1 and RW2. (A) Deformation relative to the
mean shape toward the negative direction of RW1. (B) Deformation relative to the mean
shape toward the positive direction of RW1. (C) Deformation relative to the mean shape
toward the negative direction of RW2. (D) Deformation relative to the mean shape toward
the positive direction of RW2
10.7.2 Multiple Regression analysis
Regressing the full set of partial warps on log centroid size showed no significant
differences (Wilks' Lambda = 0.4372, F (16, 13) = 1.046, P>0.4). Size explains
only 4.91% of the shape variation in our sample. Clearly, in our sample, shape is
not a function of size.
The isometric growth of appendicular skeleton in sauropods has been noted
(e.g. Carpenter and McIntosh 1994). For instance, the limbs of Camarasaurus
show evidence of isometric growth with very little indication of allometry (Wil-
hite 1999, 2003). Some authors also noted isometric growth in the limbs of Apato-
saurus (Carpenter and MacIntosh 1994).
10.8 DISCUSSION AND CONCLUSIONS
Ichnology can complement information from the dinosaur osteological record, for
instance, by providing complementary data on autopodia shape and structure as
well on limb posture. (Lockley and Hunt 1995; Gatesy et al. 1999; Wilson and
Carrano 1999).

10. Sauropod Tracks – a geometric morphometric study 137
GM techniques were applied for the first time in the Sauropodomorpha ich-
nological record, as far as we are aware. This methodology allowed to descrimi-
nate between tendencies in shape changes in our sample as well as to confirm the
absence of allometry.
The shape variation observed in our sample are caused by:
relative digit position (inward/medial to outward/lateral rotation);
medial region bending (directly associated with outward/lateral rotation of dig-
its) and relative heel position (inward/medial to outward/lateral rotation).
We have observed and quantified that specimens attributed to prosauropods
(Tetrasauropus, Pseudotetrasauropus and Paratetrasauropus) are closely related
to each other in comparison to the mean shape and to specimens attributed to
sauropods (i.e., there is a morphological discontinuity between the Prosauropoda
and Sauropoda tracks). As a consequence of this GM analysis, we can maintain
the idea of a Prosauropoda origin for Morphotype 3, which includes the above re-
ferred ichnogenus type Tetrasauropus. An opposite ichnotaxonomical proposal is
the attribution of Tetrasauropus to the Sauropoda ichnological record (Lockley et
al. 2001).
The general sauropod track record suggests that pes prints are slightly longer
than wider and present a trend as an outward rotation of four or five externally ro-
tated digit impressions (e.g. Farlow et al. 1989; Lockley 1991; Meyer et al. 1994).
These features are reliable with character 64 on the sauropod phylogenetic hy-
pothesis of Wilson and Sereno (1998) - “Pedal unguals, orientation: aligned with
(0), or deflected lateral to (1), digit axis.”.
Digit rotation is the most important factor in the shape variation in our sample.
Despite this, there are other factors contributing to the variability of shape ob-
served.
This analysis allows us to suggest that Brontopodus birdi present more outward
rotated digits than the Portuguese specimens 3, 4, 5 and 8. These specimens pre-
sent a digit rotation very similar to mean shape.
Specimens from Croatia (27, 28, 29 and 30) were attributed to a Titanosauria
origin (Dalla Vecchia and Tarlao 2000). In this analysis, they are the most extreme
specimens regarding inward digit rotation, which is close associated with Morpho-
type 3 (Prosauropoda origin) and distant to Morphotypes 1 and 2 (Sauropoda ori-
gin). This may suggest a slightest non-sauropoda origin hypothesis or digit mis-
identification (i.e., digit I could be digit IV, inverting the digit rotation course).
This methodology could permit the recognition of misidentified tracks as long as
other factors could be included (e.g. stratigraphical age, wide or narrow-gauge
trackway).
Morphotypes 1 and 2 are morphologically comparable, which is confirmed by
its parataxonomical origin affinity. This GM study corroborates the majority of
previous ichnotaxonomical classifications.
The multivariate regression analysis on the centroid size supports the lack of al-
lometry in different taxa of Sauropoda limbs (Wilhite 1999, 2003; Carpenter and
MacIntosh 1994). This geometric study corroborates the osteological results on
absence of allometry on Sauropoda appendicular structures.

138
Other multivariate analyses are currently under study using as independent
variables: velocity; wide/narrow gauge trackway; geological/stratigraphical frame;
illustration authors. This latter variable study is justified by the ichnological inter-
pretation that precedes each track illustration. It means the author of the illustra-
tion is a very important variable in geometric morphometric studies that uses track
contours.
Table 1. Reference, morphotype, parataxonomy, age, literature consulted and standard
dimensions of the specimens analyzed. * - measurements made by the authors; ** -
measurements made by the authors on the most external contour; LT- Late Triassic; J-
Jurassic MJ- Middle Jurassic UJ- Upper Jurassic; LC – Lower Cretaceous.
Reference Morphotype Parataxonomy Age Authors Length Width
1 1 Brontopodus birdi LC Thulborn 1990, p. 170, fig. 6.16.a 89* 65*
2 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 249, fig. 6.15.9. 80 58
3 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 241, fig. 6.15.5. 72* 56*
4 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 241, fig. 6.15.5. 80 50
5 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 186, fig. 6.7.3. 85 68
6 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 186, fig. 6.7.3. 70 60
7 2 Brontopodus aff. B. birdi UJ Santos 2003, p. 188, fig. 6.7.4. 60 60
8 4 Polyonichnus gomesi MJ Santos 2003, p. 124, fig. 6.1.7. 90 60
9 2 Brontopodus aff. B. birdi UJ Lires 2000, p. 33 74* 56*
10 2 Brontopodus aff. B. birdi UJ Lires 2000, p. 34 62* 58*
11 2 Brontopodus aff. B. birdi UJ Lires 2000, pers. commun. 52 36
12 1 Brontopodus sp. UJ Lockley & Mickelson 1997, p. 136 60 46
13 1 Brontopodus sp. UJ Lockley & Mickelson 1997, p. 136 55 40
14 1 Brontopodus sp. UJ Lockley & Mickelson 1997, p. 136 55 40
15 2 Brontopodus (?) LC Thulborn 1990, p. 170, fig. 6.16.f 70 65*
16 3 Pseudotetrasauropus LT Thulborn, 1990, p. 178, fig. 6.23.b 49 39*
17 3 Paratetrasauropus LT Thulborn 1990, p. 178, fig. 6.23.e 28 22*
18 3 Tetrasauropus LT Thulborn 1990, p. 178, fig. 6.23.c 44 41*
19 4 Non-identified J Thulborn 1990, p. 170, fig. 6.16.e 80 51*
20 4 Non-identified LC Woodhams 1998, pers. commun. 58* 36*
21 4 Non-identified LC Woodhams 1998, pers. commun. 62* 51*
22 1 Brontopodus birdi LC Ray Stanford 2003, pers. commun. 5.8 5
23 1 Brontopodus birdi LC Ray Stanford 2003, pers. commun. 5.8 5
24 2 Brontopodus type MJ Romano et al. 1999, p. 365, fig. 3A 94 70
25 4 Breviparopus type MJ Romano et al. 1999, p. 365, fig. 3B 81 67
26 2 Brontopodus type UJ Lockley et al. 1994, p.143, fig. 6.b 83* 51*
27 4 Titanosaurimanus nana LC Dalla Vecchia & Tarlao, 2000, p. 261, fig. 29E 34.5** 32**
28 4 Titanosaurimanus nana LC Dalla Vecchia & Tarlao, 2000, p. 261, fig. 29G 33** 29**
29 4 Titanosaurimanus nana LC Dalla Vecchia & Tarlao, 2000, p. 261, fig. 29B 31** 23**
30 4 Titanosaurimanus nana LC Dalla Vecchia & Tarlao, 2000, p. 261, fig. 29F 34** 29**

10. Sauropod Tracks – a geometric morphometric study 139

10.9 Acknowledgements
The authors would like to thank: Prof. James Rohlf (Stony Brook University,
USA) for all of his helpful comments and support; Dr. Ângela Delgado Buscalioni
and Jesus Marugán-Lóbon (Universidad Autónoma, Madrid) for helpful com-
ments; Jose Lires (University of Oviedo, Spain) for giving information about
sauropod pes natural casts of Upper Jurassic of Asturias, Spain; Ray Stanford
(Mesozoic Track Project) for the two small Brontopodus birdi specimens (22, 23)
and all of his helpful comments; Dr. Kenneth Woodhams for the specimens 20 and
21; Sara Bárrios for assistance in data processing.
A special thanks to Prof. Galopim de Carvalho (MNHN/University of Lisbon) for
all of his support.
The manuscript was greatly improved by the comments of two anonymous re-
viewers.
This work was partially funded by Project Fundação para a Ciência e Tecnologia
(FCT) POCTI/PAL/36550/2000 - “Dinosaur Osteological and Ichnological studies
of the Mesozoic of Portugal (DINOS)”.
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Table of Contents
1 Introduction ........................................................................................................1
Ashraf M. T. Elewa ...........................................................................................1
References .....................................................................................................4
2 Application of geometric morphometrics to the study of shape polymor-
phism in Eocene ostracodes from Egypt and Spain............................................7
Ashraf M. T. Elewa ...........................................................................................7
2.1 Abstract ...................................................................................................7
2.2 Introduction .............................................................................................7
2.3 Brief notes on morphometrics .................................................................9
2.4 Polymorphism in ostracodes..................................................................10
2.5 Materials and methods...........................................................................11
2.6 Results ...................................................................................................14
2.6.1 The Egyptian material....................................................................14
2.6.2 The Spanish material......................................................................19
2.7 Conclusions ...........................................................................................24
2.8 Acknowledgements ...............................................................................26
References ...................................................................................................26
3 Morphometric analysis of population differentiation and sexual
dimorphism in the blue spiny lobster Panulirus inflatus (Bouvier 1895)
from NW Mexico .................................................................................................29
Francisco Javier García-Rodríguez, José de la Cruz Agüero, Ricardo
Pérez-Enriquez and Norman MacLeod............................................................29
3.1 Abstract .................................................................................................29
3.2 Introduction ...........................................................................................29
3.3 Material and methods ............................................................................31
3.4 Results ...................................................................................................33
3.5 Discussion .............................................................................................37
3.6 Acknowledgements ...............................................................................40
References ...................................................................................................40
4. The effect of alcohol and freezing preservation on carapace size and
shape in Liocarcinus depurator (Crustacea, Brachyura) ................................45
Marta Rufino, Pere Abelló and Andrew B. Yule .............................................45
4.1 Abstract .................................................................................................45
4.2 Introduction ...........................................................................................45

X Table of Contents
4.3 Materials and methods .......................................................................... 47
4.4 Results................................................................................................... 48
4.5 Discussion ............................................................................................. 51
4.6 Acknowledgements ............................................................................... 52
References................................................................................................... 52
5 Allometric field decomposition – an attempt at morphogenetic
morphometrics..................................................................................................... 55
Øyvind Hammer .............................................................................................. 55
5.1 Abstract ................................................................................................. 55
5.2 Introduction........................................................................................... 55
5.3 Allometric fields.................................................................................... 56
5.4 Allometric field decomposition............................................................. 61
5.5 Case study: Ammonite allometry .......................................................... 62
5.6 Conclusion ............................................................................................ 64
References................................................................................................... 65
6 A combined landmark and outline-based approach to ontogenetic shape
change in the Ordovician trilobite Triarthrus becki ........................................ 67
H. David Sheets, Keonho Kim and Charles E. Mitchell.................................. 67
6.1 Abstract ................................................................................................. 67
6.2 Introduction........................................................................................... 68
6.3 Materials................................................................................................ 70
6.4 Methods................................................................................................. 71
6.5 Results................................................................................................... 76
6.6 Concerns about the use of semi-landmarks ........................................... 80
6.7 Acknowledgements ............................................................................... 81
References................................................................................................... 81
7 Morphological analysis of two- and three-dimensional images of
branching sponges and corals ............................................................................ 83
Jaap A. Kaandorp and Rafael A. Garcia Leiva ................................................ 83
7.1 Abstract ................................................................................................. 83
7.2 Introduction........................................................................................... 83
7.3 Methods................................................................................................. 87
7.3.1 Measurements in two-dimensional images .................................... 87
7.3.2 Three-dimensional data acquisition ............................................... 90
7.3.3 Three-dimensional measurements based on the
morphological skeleton.......................................................................... 90
7.4 Results................................................................................................... 92
7.5 Discussion ............................................................................................. 92
7.6. Acknowledgements .............................................................................. 94
References................................................................................................... 94

Table of Contents XI
8 Geometric morphometric analysis of head shape variation in four
species of hammerhead sharks (Carcharhiniformes: Sphyrnidae).................97
Mauro J. Cavalcanti .........................................................................................97
8.1 Abstract .................................................................................................97
8.2 Introduction ...........................................................................................98
8.3 Materials and methods...........................................................................99
8.3.1 Samples..........................................................................................99
8.3.2 Data acquisition ...........................................................................100
8.3.3 Data analysis ................................................................................100
8.4 Results .................................................................................................102
8.5 Discussion ...........................................................................................110
8.6 Acknowledgements .............................................................................111
References .................................................................................................111
9 Morphometric stock structure of the Pacific sardine Sardinops sagax
(Jenyns, 1842) off Baja California, Mexico .....................................................115
José De La Cruz Agüero and Francisco Javier García Rodríguez .................115
9.1 Abstract ...............................................................................................115
9.2 Introduction. ........................................................................................116
9.3 Materials and methods.........................................................................117
9.3.1 Sample collection and treatment of data ......................................117
9.32 Data analysis .................................................................................119
9.4 Results .................................................................................................120
9.4.1 Data improvement........................................................................120
9.4.2 Univariate analysis.......................................................................120
9.4.3 Multivariate analysis....................................................................121
9.5 Discussion ...........................................................................................122
9.6 Acknowledgements .............................................................................124
References .................................................................................................125
10. Sauropod Tracks – a geometric morphometric study ..............................129
Luis Azevedo Rodrigues and Vanda Faria dos Santos ..................................129
10.1 Abstract .............................................................................................129
10.2 Introduction .......................................................................................129
10.3 Materials and methods.......................................................................130
10.3.1 Samples......................................................................................130
10.3.2 Obtaining landmarks coordinates...............................................131
10.3.3 Description of landmarks...........................................................132
10.4 Relative warp analysis.......................................................................133
10.5 Multiple regression analysis ..............................................................133
10.6 Software ............................................................................................134
10.7 Results ...............................................................................................134
10.7.1 Relative warps analysis..............................................................134
10.7.2 Multiple regression analysis ......................................................136
10.8 Discussion and conclusions...............................................................136

XII Table of Contents
10.9 Acknowledgements ........................................................................... 139
References................................................................................................. 139
11 Morphometric approach to Titanosauriformes (Sauropoda, Dinosauria)
femora: Implications to the paleobiogeographic analysis .............................. 143
José I. Canudo and Gloria Cuenca-Bescós .................................................... 143
11.1 Abstract ............................................................................................. 143
11.2 Introduction....................................................................................... 143
11.3 Materials and methods ...................................................................... 146
11.4 Results and discussions ..................................................................... 149
11.4.1 Titanosauriformes of the Lower Cretaceous.............................. 149
11.4.2 Titanosauria and Titanosauridae ................................................ 150
11.4.3 Titanosauria of Laurasia ............................................................ 151
11.4.4 Titanosauria of Gondwana of Upper Cretaceous. Alamosaurus,
the emigrant ............................................................................... 152
11.5 Conclusions ....................................................................................... 153
11.6 Acknowledgements ........................................................................... 154
References................................................................................................. 154
12 Geometric morphometrics in macroevolution: morphological diversity of
the skull in modern avian forms in contrast to some theropod dinosaurs ... 157
Jesús Marugán-Lobón and Ángela D. Buscalioni.......................................... 157
12.1 Abstract ............................................................................................. 157
12.2 Introduction....................................................................................... 158
12.2.1 Theoretical perspective .............................................................. 158
12.2.2 Morphology ............................................................................... 159
12.2.3 Phylogenetic context.................................................................. 160
12.3 Materials and methods ...................................................................... 161
12.4 Results............................................................................................... 162
12.5. Discussion and conclusions.............................................................. 168
12.6 Acknowledgements ........................................................................... 171
References................................................................................................. 171
13 Correlation of foot sole morphology with locomotion behaviour and
substrate use in four passerine genera............................................................. 175
Fränzi Korner-Nievergelt............................................................................... 175
13.1 Abstract ............................................................................................. 175
13.2 Introduction....................................................................................... 175
13.3. Species and data ............................................................................... 176
13.3.1 Species and sample size............................................................. 176
13.3.2 Morphological data .................................................................... 177
13.2.3 Behavioural data ........................................................................ 178
13.2.4 Statistics..................................................................................... 180
13.3 Results............................................................................................... 183
13.4 Discussion ......................................................................................... 187
13.4.1 Reconstruction of mean foot sole shapes................................... 187

Table of Contents XIII
13.4.2 Parallelism .................................................................................188
13.4.3 Functional aspects of plantar morphological traits ....................189
13.5 Acknowledgements ...........................................................................191
References .................................................................................................192
Appendix ...................................................................................................195
Mean ecological scores.........................................................................195
14 Maximum-likelihood identification of fossils: taxonomic identification of
Quaternary marmots (Rodentia, Mammalia) and identification of vertebral
position in the pipesnake Cylindrophis (Serpentes, Reptilia) ........................197
P. David Polly and Jason J. Head ..................................................................197
14.1 Abstract .............................................................................................197
14.2 Introduction .......................................................................................198
14.3 Materials and methods.......................................................................200
14.3.1 Marmots .....................................................................................200
14.3.2 Snakes ........................................................................................203
14.3.3 ML identification procedure ......................................................205
14.3.4 Cross-validation assessment ......................................................206
14.3.5 Identification of unknowns ........................................................207
14.4 Results ...............................................................................................207
14.4.1 Marmots .....................................................................................207
14.4.2 Snakes ........................................................................................211
14.5 Discussion .........................................................................................213
14.6 Conclusions .......................................................................................217
14.7 Acknowledgements ...........................................................................218
References .................................................................................................218
15 Geometric morphometrics of the upper antemolar row configuration
in the brown-toothed shrews of the genus Sorex (Mammalia) ......................223
Igor Y. Pavlinov.............................................................................................223
15.1 Abstract .............................................................................................223
15.2 Introduction .......................................................................................223
15.3 Materials and methods.......................................................................225
15.4 Results ...............................................................................................226
15.5 Conclusions .......................................................................................229
15.6 Acknowledgements ...........................................................................230
References .................................................................................................230
16 Geometric morphometrics in paleoanthropology: Mandibular shape varia-
tion, allometry, and the evolution of modern human skull morphology ......231
Markus Bastir and Antonio Rosas .................................................................231
16.1 Abstract .............................................................................................231
16.2 Introduction .......................................................................................231
16.2 Material and methods ........................................................................234
16.3 Geometric morphometry ...................................................................234
16.3.1 Thin-plate splines.......................................................................235

XIV Table of Contents
16.3.2 Missing data............................................................................... 235
16.3.3 Geometric morphometric software and data analyses ............... 236
16.4 Results............................................................................................... 236
19.5. Discussion ........................................................................................ 238
19.7 Conclusions ....................................................................................... 240
16.9. Acknowledgements .......................................................................... 241
References................................................................................................. 241
17 3-D geometric morphometric analysis of temporal bone landmarks in
Neanderthals and modern humans.................................................................. 245
Katerina Harvati ............................................................................................ 245
17.1 Abstract ............................................................................................. 245
17.2 Introduction....................................................................................... 245
17.3 Materials and methods ...................................................................... 246
17.4 Results............................................................................................... 248
17.5 Discussion ......................................................................................... 253
17.5.1 Modern humans ......................................................................... 253
17.5.2 Neanderthals .............................................................................. 253
17.5.3 Upper Paleolithic Europeans ..................................................... 254
17.5.4 Kabwe........................................................................................ 255
17.6 Conclusions ....................................................................................... 256
17.7 Acknowledgements ........................................................................... 256
References................................................................................................. 256

Index .................................................................................................................. 259