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Zoological Journal of the Linnean Society, 2019, 186, 1006–1025. With 8 figures.
© 2019 The Linnean Society of London, Zoological Journal of the Linnean Society, 2019, 186, 1006–1025
Comparative morphology of presacral vertebrae in
extant crocodylians: taxonomic, functional and ecological
implications
MASAYA IIJIMA
1,2,
* and TAI KUBO
3
1
School of Resources and Environmental Engineering, Hefei University of Technology, Hefei, Anhui
230009, China
2
Department of Natural History Sciences, Hokkaido University, N10W8 Kita-ku, Sapporo, Hokkaido
0600810, Japan
3
The University Museum, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 1130033, Japan
Received 27 June 2018; revised 22 November 2018; accepted for publication 25 December 2018
Despite its systematic and functional relevance, the axial skeleton of crocodylians has received considerably less
attention than the cranial and appendicular skeleton due to the assumed evolutionary conservativeness. The
current study provides comprehensive comparisons of presacral vertebrae in extant crocodylians to demonstrate:
(1) taxonomic variation, (2) size-dependent shape changes and (3) potential integration between vertebrae and skull
functional modules. Our comparisons highlighted the uniqueness of the Indian gharial, Gavialis gangeticus, among
extant crocodylians. The presacral vertebrae of G. gangeticus are characterized by reduced level of regionalization and
increased intervertebral mobility in the neck (more narrowly placed zygapophyses and short vertebral processes),
which would help lateral head sweeping under water for foraging. The scaling relationships of vertebral dimensions
against the body size proxy indicate that dorsal vertebrae become stiffer with increasing body size: positive
allometries were observed in areas and heights of inter-central joints, and lengths of vertebral processes (neural
spines and transverse processes). These structural changes presumably serve to resist increasing compression
loads and dorsiflexion bending moment on dorsal vertebrae of larger individuals during terrestrial locomotion. The
analyses of correlation between vertebral dimensions and snout shape revealed that slender-snouted species tend to
have more laterally mobile necks (specifically narrow zygapophyses), implying the potential integration of cranial
and neck modules to optimize the foraging strategy.
KEYWORDS: aquatic adaptation – Crocodylia – postcranial skeleton – ontogeny.
INTRODUCTION
Crocodylians are the largest known living reptiles and
at least 27 species are distributed in the tropics and
subtropics around the world (Grigg & Kirshner, 2015).
As derived members of Pseudosuchia, a diverse and
successful group of reptiles that rivalled early dinosaurs
in the Triassic (Brusatte et al., 2008), crocodylians
have survived several large environmental crises
(Brochu, 2003).
Recent years have seen a rise in the number of
comparative studies on postcrania of extant and fossil
crocodylians. Such studies often focused on girdle
and limb elements, revealing the morphofunctional
differences between the two large groups, Alligatoridae
and Crocodylidae. Compared to crocodylids,
alligatorids are characterized by a stouter humerus
and coracoid, shorter stylopodia (humerus and femur)
in fore- and hindlimbs, shorter muscle fascicle lengths
and larger cross-sectional areas for pectoral limb
muscles, potentially reflecting the differences in their
locomotor ecology (Chamero et al., 2013; Allen et al.,
2015; Iijima et al., 2018). However, the axial skeleton
of extant crocodylians has received considerably
less attention, despite its potential taxonomic and
ecological significance. This is partly because vertebral
osteology and myology of extant crocodylian families
were regarded as remarkably conservative and similar
to each other (Hoffstetter & Gasc, 1969). Consequently, *Corresponding author. E-mail: miijima8@gmail.com
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