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Background
Little is known about the long-term patterns of body size
evolution in Crocodylomorpha, the >â200-million-year-old
group that includes living crocodylians and their extinct
relatives. Extant crocodylians are mostly large-bodied
(3â7âm) predators. However, extinct crocodylomorphs exhibit
a wider range of phenotypes, and many of the earliest taxa
were much smaller (<â1.2âm). This suggests a pattern of
size increase through time that could be caused by
multi-lineage evolutionary trends of size increase or by
selective extinction of small-bodied species. Here, we
characterise patterns of crocodylomorph body size evolution
using a model fitting-approach (with cranial measurements
serving as proxies). We also estimate body size disparity
through time and quantitatively test hypotheses of biotic
and abiotic factors as potential drivers of crocodylomorph
body size evolution.
Results
Crocodylomorphs reached an early peak in body size disparity
during the Late Jurassic, and underwent an essentially
continual decline since then. A multi-peak
Ornstein-Uhlenbeck model outperforms all other evolutionary
models fitted to our data (including both uniform and
non-uniform), indicating that the macroevolutionary dynamics
of crocodylomorph body size are better described within the
concept of an adaptive landscape, with most body size
variation emerging after shifts to new macroevolutionary
regimes (analogous to adaptive zones). We did not find
support for a consistent evolutionary trend towards larger
sizes among lineages (i.e., Copeâs rule), or strong
correlations of body size with climate. Instead, the
intermediate to large body sizes of some crocodylomorphs are
better explained by group-specific adaptations. In
particular, the evolution of a more aquatic lifestyle
(especially marine) correlates with increases in average
body size, though not without exceptions.
Conclusions
Shifts between macroevolutionary regimes provide a better
explanation of crocodylomorph body size evolution on large
phylogenetic and temporal scales, suggesting a central role
for lineage-specific adaptations rather than climatic
forcing. Shifts leading to larger body sizes occurred in
most aquatic and semi-aquatic groups. This, combined with
extinctions of groups occupying smaller body size regimes
(particularly during the Late Cretaceous and Cenozoic), gave
rise to the upward-shifted body size distribution of extant
crocodylomorphs compared to their smaller-bodied terrestrial
ancestors.