Name: Emma Reinke

Student #: 1007113687
TA: Alejandro Lopez

Jaw joint complexity in crocodilians has constrained cranial morphology which has led to a
decline in disparity

Introduction
The evolution of cranial complexity in crocodilians has evolved to make them optimal
hunters. Crocodiles have the largest jaw force of any animal on the planet around 3700 psi
((Erickson 2018)). This awesome display of power is the compilation of hundreds of millions of
years of evolution. This powerful jaw force is believed to be an adaptation from the age of the
dinosaurs, when modern day crocodiles much larger ancestors had to compete with gigantic
dinosaurs with impressive bite forces in the Mesozoic era ((Federation of American Societies for
Experimental Biology (FASEB) 2016 Apr 4)).
Crocodylians have a special second joint in their jaw that helps distribute jaw force
throughout the skull. This second jaw joint was found by a research team led by Casey Holliday
et al. They used 3-D modelling tools to map the bones and cartilage in the crocodylians skull
((Federation of American Societies for Experimental Biology (FASEB) 2016 Apr 4)). In this
joint is a specialized muscle in their neck called the ventral pterygoideus muscle which gives the
gives the joint the power in needs to have such a large jaw force ((Gignac and O’Brien 2016)).
This specialized jaw joint has allowed crocodylians to dominate in their ecological niche for
millions of years. Although, considering how long crocodiles have been around for there appears
to be a startlingly low amount of disparity in the crocodylian order compared to their sister clade,
birds.
Crocodylians are part of a subclass called Archosaurs, a group that includes crocodylians,
birds, and the now extinct dinosaurs. Crocodylians and birds diverged from their MRCA around
240 MYA ((Green et al. 2014)), although crocodylians appear to have a slower rate of evolution
compared to their sister taxa, birds, this can be seen in figure 1. The bird phylogenetic tree has a
greater disparity than crocodylians. This vast range in disparity is believed to be because of the
jaw joint complexity in crocodilians which has constrained cranial morphology. If this is the
case, then we should expect to see a decline in variance of crocodylian skull morphology
compared to bird morphology. Additionally, if this is not the case then we would expect to find
that the variance in skull morphology has remained steady and that there is no significant change
in variance and that the level of variance is similar to that of birds.

Methods
To determine whether the lower disparity in crocodylians is because of cranial
morphology a measure of the levels of variation between extinct and extant crocodiles will be
conducted. Intact skulls from Pre-Jurassic will be scanned and compared to that of extant
crocodylians. The independent variable will be time and the dependent variable will be the size
of the pre-Jurassic and extant jaw joints. This will show the variance of jaw joint size over time.
To measure the size of the joints a scanner will create a 3-D model to accurately get a reading on
the precise measurements of the size of the joints by looking at their origin and their insertion
and using a complex system to create an accurate measurement for the size of the joint. This
same test will be done with extinct birds and extant birds to show the levels of jaw joint variance
in birds which can then be used a comparison to crocodylians in the next test.
A comparison of within clade jaw joint variation between crocodylians and birds will be
done to compare the crocodylians to another clade in the archosaurs that have been
independently evolving for hundreds of millions of years. The independent variable for the
crocodylian comparison will be number of crocodylian species and the dependent variable will
be the size of the jaw joints. In the second comparison the independent variable will be the
number of bird species and the dependent variable will be the size of the jaw joints. These
distributions in jaw formation will tell whether crocodylians have a low variance compared to
birds.

Predictions
If jaw joint variation is lower in crocodylians than that of birds, then the experiment will
fail to reject the hypothesis and reject the alternative hypothesis. If there is no noticeable
difference in variation levels between crocodylians and bird or that there is less disparity in birds
than crocodylians then the hypothesis will be rejected, and the alternative hypothesis will be
supported.
Discussion
If it is found that jaw joint disparity has declined in crocodylians since their ancestors,
then this will help support the hypothesis and reject the alternative hypothesis. This could mean
that after the ancestors went extinct from their wider niche, the crocodylians that survived had
jaw joints that were so specialized that they could not adapt to fill those niches. This
specialization allows them greater jaw force and helps them rule their ecological habitats, but
they cannot adapt their jaw formations without it being detrimental to their survival. So, they are
‘stuck’ with limited phenotypes because they are limited in their ability to shift their cranial
structures (Felice et al. 2021). With their specialized jaw force the crocodylian have a much
greater range of predation which could also correspond to why they don’t seem to have evolved
in their jaw formations (Villamarín et al. 2018).
If it is found that the disparity is equal to or more than birds and greater than their
ancestors, the hypothesis is rejected and the alternative hypothesis is supported. This would
mean that there is some other reason for the limited families in crocodylia. It has been proposed
that one of the reasons that crocodylians did not expand into their ancestors’ niches was because
terrestrial land animals may have evolved at a faster rate and taken that niche before the
crocodylians were able to (Stubbs et al. 2021). Additionally, another reason other than joint
specificity for crocodylians low variation may be because of snout formation which has been a
major topic of discussion the past couple decades (Felice et al. 2021). These ideas could be an
area of further research that could be explored in future experiments.
Figures

Figure 1- A phylogenetic diagram of the evolutionary

history of mammals, crocodiles, and birds
(Image credit: Richard E. Green et al.)

Literature Cited

Deeming DC, Harrison SL, Sutton GP. 2022. Inter‐relationships among body mass, jaw
musculature and bite force in birds. Journal of Zoology. 317(2):129–137. doi:10.1111/jzo.12966.
[accessed 2022 Nov 24].
https://zslpublications.onlinelibrary.wiley.com/doi/epdf/10.1111/jzo.12966?src=getftr.

Erickson GM. 2018. The Baddest Bite. Scientific American. 318(3):40–45.

doi:10.1038/scientificamerican0318-40. [accessed 2022 Nov 17].
https://www.scientificamerican.com/article/how-crocs-came-to-rule-the-water-rsquo-s-edge/.
Federation of American Societies for Experimental Biology (FASEB). 2016 Apr 4. Scientists
find surprise lurking in crocodilian jaw. ScienceDaily. [accessed 2022 Nov 22].
https://www.sciencedaily.com/releases/2016/04/160404221029.htm.

Felice RN, Pol D, Goswami A. 2021. Complex macroevolutionary dynamics underly the
evolution of the crocodyliform skull. Proceedings of the Royal Society B: Biological Sciences.
288(1954):20210919. doi:10.1098/rspb.2021.0919. [accessed 2022 Nov 24].
https://royalsocietypublishing.org/doi/full/10.1098/rspb.2021.0919.

Felice RN, Tobias JA, Pigot AL, Goswami A. 2019. Dietary niche and the evolution of cranial
morphology in birds. Proceedings of the Royal Society B: Biological Sciences.
286(1897):20182677. doi:10.1098/rspb.2018.2677. https://doi.org/10.1098/rspb.2018.2677.

Gignac P, O’Brien H. 2016. Suchian Feeding Success at the Interface of Ontogeny and
Macroevolution. Integrative and Comparative Biology. 56(3):449–458. doi:10.1093/icb/icw041.
[accessed 2022 Nov 21]. https://doi.org/10.1093/icb/icw041.

Green RE, Braun EL, Armstrong J, Earl D, Nguyen N, Hickey G, Vandewege MW, St John JA,
Capella-Gutiérrez S, Castoe TA, et al. 2014. Three crocodilian genomes reveal ancestral patterns
of evolution among archosaurs. Science (New York, NY). 346(6215):1254449.
doi:10.1126/science.1254449. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4386873/#R5.

Holliday CM. 2013. Form, function, and evolution of archosaur mandibular symphyses. The
FASEB Journal. 27(S1). doi:10.1096/fasebj.27.1_supplement.79.6.

Stubbs TL, Pierce SE, Elsler A, Anderson PSL, Rayfield EJ, Benton MJ. 2021. Ecological
opportunity and the rise and fall of crocodylomorph evolutionary innovation. Proceedings of the
Royal Society B: Biological Sciences. 288(1947). doi:10.1098/rspb.2021.0069. [accessed 2021
May 7]. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8059953/.

Villamarín F, Jardine TD, Bunn SE, Marioni B, Magnusson WE. 2018. Body size is more
important than diet in determining stable-isotope estimates of trophic position in crocodilians.
Scientific Reports. 8(1). doi:10.1038/s41598-018-19918-6. [accessed 2022 Nov 24].
https://www.nature.com/articles/s41598-018-19918-6.