Cited by 218Paleontology and Evolutionary BiologyEvolution and Paleontology StudiesComparative Animal Anatomy Studies

The DOI matches its Crossref record; 54 of the 54 references it cites were found in a registry or an index; 3 of the 42 sources its claims cite were checked for support.

Insights into the Ecology and Evolutionary Success of Crocodilians Revealed through Bite-Force and Tooth-Pressure Experimentation

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Abstract

BACKGROUND: Crocodilians have dominated predatory niches at the water-land interface for over 85 million years. Like their ancestors, living species show substantial variation in their jaw proportions, dental form and body size. These differences are often assumed to reflect anatomical specialization related to feeding and niche occupation, but quantified data are scant. How these factors relate to biomechanical performance during feeding and their relevance to crocodilian evolutionary success are not known. METHODOLOGY/PRINCIPAL FINDINGS: We measured adult bite forces and tooth pressures in all 23 extant crocodilian species and analyzed the results in ecological and phylogenetic contexts. We demonstrate that these reptiles generate the highest bite forces and tooth pressures known for any living animals. Bite forces strongly correlate with body size, and size changes are a major mechanism of feeding evolution in this group. Jaw shape demonstrates surprisingly little correlation to bite force and pressures. Bite forces can now be predicted in fossil crocodilians using the regression equations generated in this research. CONCLUSIONS/SIGNIFICANCE: Critical to crocodilian long-term success was the evolution of a high bite-force generating musculo-skeletal architecture. Once achieved, the relative force capacities of this system went essentially unmodified throughout subsequent diversification. Rampant changes in body size and concurrent changes in bite force served as a mechanism to allow access to differing prey types and sizes. Further access to the diversity of near-shore prey was gained primarily through changes in tooth pressure via the evolution of dental form and distributions of the teeth within the jaws. Rostral proportions changed substantially throughout crocodilian evolution, but not in correspondence with bite forces. The biomechanical and ecological ramifications of such changes need further examination.

Overview

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TL;DR

Erickson et al. measured bite forces and tooth pressures across all 23 extant crocodilian species (83 adults) and found that body mass, not rostral shape, explains nearly all bite-force variation. Crocodilians generate the highest bite forces and tooth pressures known among living animals, with tooth pressures exceeding the shear strength of bone and previously reported maxima.

Key findings

  • Molariform bite forces ranged from 900 to 8,983 N (*Paleosuchus palpebrosus* and *Crocodylus porosus*, respectively); one *C. porosus* individual reached 16,414 N, the highest bite force measured in any animal, exceeding the 4,500 N recorded for spotted hyena.
  • Body mass was the primary determinant of bite force (TIPs: R² = 0.92; PIC: R² = 0.87, p < 0.0001), with a scaling coefficient of 0.708 ± 0.111, not statistically different from isometry (0.667). Rostral proportions explained just 19% of remaining size-standardized phylogenetically corrected variance (2.5% of total variance; p = 0.03).
  • Caniniform tooth pressures ranged from 195 to 1,344 MPa; molariform pressures ranged from 203 to 1,388 MPa. All exceeded the 147 MPa reported for *Dunkleosteus*, some by as much as 17-fold, and all exceeded the 65–71 MPa ultimate shear strength of bone.
  • Tooth pressures scaled positively allometric (caniniform: 0.490 ± 0.203; molariform: 0.553 ± 0.180 vs. isometry 0.000) and were uncorrelated with rostral proportions (PIC R² = 0.001 caniniform; 0.094 molariform). *Gavialis gangeticus* was the only statistical low-force outlier; fossil predictions included 27,531–34,424 N for 6.7 m *C. porosus* and a historical range of 628–102,803 N.

Methods

  • 83 sexually mature adult specimens representing all 23 extant crocodilian species were tested from St. Augustine Alligator Farm Zoological Park and Crocodylus Park; size range was 1.24–4.59 m and 7–531 kg, with multiple individuals for 19 species.
  • Bite forces were recorded using sandwich transducers and a portable charge amplification system; 3–5 bites were recorded per animal and the highest used. Forces were measured at the prominent maxillary molariform tooth, and caniniform forces were inferred by moment calculation. Tooth contact areas came from dental putty molds and epoxy casts indented 1 mm into clay, measured with ImageJ; pressures were force divided by contact area.
  • Analyses used OLS and reduced major axis regressions on log-transformed taxon representative data, raw “TIPs” analyses, and phylogenetically independent contrasts (PIC) based on a re-estimated DNA sequence phylogeny from Gatesy et al. data; K statistics and ancestral-state reconstructions were also used, with statistical significance reported for PICs.

Limitations

  • Stated: competing phylogenetic hypotheses exist and some sub-clade relationships are debated; the tree used is the best available estimate, and exploring other hypotheses was beyond scope. The upper-bound bite-force estimate for *Deinosuchus riograndensis* is more tenuous because the largest fossil specimens greatly exceed the neontological size range studied.
  • Stated: caniniform pressure estimation was not possible for taxa with shed or broken caniniform teeth; initial tooth pressures alone are insufficient to single out durophagous species’ biomechanical import, and rostral shape needs further explanation.
  • Evident from the text and tables: sample sizes are highly uneven, with many species represented by N = 1 (e.g., *Crocodylus intermedius*, *C. mindorensis*, *C. moreletii*, *C. palustris*), and all specimens were captive, conservation, or display animals from two facilities, so wild population variation is not directly sampled, although the paper cites prior captive–wild equivalence for *Alligator mississippiensis*.

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Integrity signals

Registry

Crossref · 10.1371/journal.pone.0031781

Cites

54 of 54 references verified0 citation errors · 0 not checkable

Statistics

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Figures

Figure 1 Phylogenetic hypothesis for extant Crocodylia showing variation in rostral proportions. The cladogram is based on reanalysis (see Materials and Methods) of molecular data from Gatesy and colleagues [2] using maximum likelihood and non-parametric rate-smoothing with branch lengths proportional to time. Lineages shown in blue represent caiman (a–e) and alligators (f,g) ( = Alligatoridae), and those in green crocodiles (h–t) and gharials (u,v) ( = Crocodylidae+Gavialidae). The Yacare caiman, Caiman yacare is not shown for it was not utilized in the Gatesy et al. [2] analysis. Dorsal views of heads are modified from Wermuth and Fuchs [53] and standardized to the same length to show relative differences in rostral form. Bracketed numbers following taxon names are the mean rostral proportions or RP ( = mid-rostral width/snout length) for each taxon from our study. Phylogenetic Independent Contrasts were performed on these 22 species; however, bite force, tooth pressure, and morphometric measurements and subsequent TIPs analyses were performed for all 23 extant taxa, including Caiman yacare.
Figure 1 Phylogenetic hypothesis for extant Crocodylia showing variation in rostral proportions. The cladogram is based on reanalysis (see Materials and Methods) of molecular data from Gatesy and colleagues [2] using maximum likelihood and non-parametric rate-smoothing with branch lengths proportional to time. Lineages shown in blue represent caiman (a–e) and alligators (f,g) ( = Alligatoridae), and those in green crocodiles (h–t) and gharials (u,v) ( = Crocodylidae+Gavialidae). The Yacare caiman, Caiman yacare is not shown for it was not utilized in the Gatesy et al. [2] analysis. Dorsal views of heads are modified from Wermuth and Fuchs [53] and standardized to the same length to show relative differences in rostral form. Bracketed numbers following taxon names are the mean rostral proportions or RP ( = mid-rostral width/snout length) for each taxon from our study. Phylogenetic Independent Contrasts were performed on these 22 species; however, bite force, tooth pressure, and morphometric measurements and subsequent TIPs analyses were performed for all 23 extant taxa, including Caiman yacare.CC BY · PubMed Central
Figure 2 Skull and jaws of a wild adult American alligator (Alligator mississippiensis) showing the prominent teeth used for initially seizing and crushing prey. The most prominent caniniform and molariform teeth of the upper jaw that are associated with the convexities in the maxilla are highlighted. Because of their greater length relative to the adjacent teeth, and the propensity of crocodilians to bite unilaterally, the crowns of these teeth typically initiate contact with prey during biting. This specimen demonstrates the natural in situ condition of the teeth, which often fall out during skeletonization and must be reattached. As such the natural prominence of the teeth is sometimes not represented in prepared specimens. The caniniform teeth in crocodilians are longer, more slender, and generally have rounder cross-sectional shapes than the molariform teeth. In A. mississippiensis the apices of the caniniforms are fairly dull, whereas in more piscivorous species they are sharp and needle-like. Besides being utilized for seizing prey, caniniform teeth are also used in fighting, defense, aggression, and display. Crocodilian molariforms, on the other hand, are shorter and are typically blunter-tipped than the caniniform teeth. They range interspecifically from having a rounded bulbous morphology to being laterally compressed and blade-like. The intermediate condition seen here is characteristic of A. mississippiensis. Molariform teeth are primarily used for crushing and gripping prey in preparation for swallowing, but are also utilized for display and seizing prey.
Figure 2 Skull and jaws of a wild adult American alligator (Alligator mississippiensis) showing the prominent teeth used for initially seizing and crushing prey. The most prominent caniniform and molariform teeth of the upper jaw that are associated with the convexities in the maxilla are highlighted. Because of their greater length relative to the adjacent teeth, and the propensity of crocodilians to bite unilaterally, the crowns of these teeth typically initiate contact with prey during biting. This specimen demonstrates the natural in situ condition of the teeth, which often fall out during skeletonization and must be reattached. As such the natural prominence of the teeth is sometimes not represented in prepared specimens. The caniniform teeth in crocodilians are longer, more slender, and generally have rounder cross-sectional shapes than the molariform teeth. In A. mississippiensis the apices of the caniniforms are fairly dull, whereas in more piscivorous species they are sharp and needle-like. Besides being utilized for seizing prey, caniniform teeth are also used in fighting, defense, aggression, and display. Crocodilian molariforms, on the other hand, are shorter and are typically blunter-tipped than the caniniform teeth. They range interspecifically from having a rounded bulbous morphology to being laterally compressed and blade-like. The intermediate condition seen here is characteristic of A. mississippiensis. Molariform teeth are primarily used for crushing and gripping prey in preparation for swallowing, but are also utilized for display and seizing prey.CC BY · PubMed Central
Figure 3 Taxon representative adult bite forces for extant Crocodylia with respect to mean body mass and the relationship between rostral proportion and force generation. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the strong correspondence between body mass and bite force. Extant alligators and caiman (Alligatoridae), and crocodiles (Crocodylidae) show comparable relative bite-force capacities. Note that only Gavialis gangeticus is a statistical (i.e. outside the 95% confidence interval) low-force outlier. (B) Linear regression of the size-standardized residual bite force versus rostral proportion phylogenetic independent contrasts showing the low correlation between these after accounting for phylogeny and body mass.
Figure 3 Taxon representative adult bite forces for extant Crocodylia with respect to mean body mass and the relationship between rostral proportion and force generation. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the strong correspondence between body mass and bite force. Extant alligators and caiman (Alligatoridae), and crocodiles (Crocodylidae) show comparable relative bite-force capacities. Note that only Gavialis gangeticus is a statistical (i.e. outside the 95% confidence interval) low-force outlier. (B) Linear regression of the size-standardized residual bite force versus rostral proportion phylogenetic independent contrasts showing the low correlation between these after accounting for phylogeny and body mass.CC BY · PubMed Central
Figure 4 Caniniform pressure values for extant Crocodylia, their phylogenetic distribution, and inferred ancestral character states. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the weak relationship between body mass and caniniform pressure. Note that slender-snouted piscivorous to semi-piscivorous ecomorphs (Gavialis gangeticus and Crocodylus intermedius, respectively) show exceptionally high-pressure values (outside the 95% confidence interval), and Crocodylus johnsoni shows pressures expected of animals nearly a magnitude in size larger. Other ecomorphs show much lower and similar relative values. The arrow indicates the typical ultimate shear strength of bone. (B) Ancestral character-state reconstruction using squared-change parsimony of size-standardized caniniform pressures. Residual caniniform pressure values are color coded to MPa (squared-change parsimony; squared length = 19.491). Vertical scale is in relative time, with the outgroup/ingroup root arbitrarily set to 1.0. High relative pressures were achieved independently in Crocodylus intermedius, Gavialis gangeticus, and Crocodylus johnsoni. Uncolored branches represent taxa for which the caniniform teeth were shed or broken, and so pressure estimation was not possible.
Figure 4 Caniniform pressure values for extant Crocodylia, their phylogenetic distribution, and inferred ancestral character states. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the weak relationship between body mass and caniniform pressure. Note that slender-snouted piscivorous to semi-piscivorous ecomorphs (Gavialis gangeticus and Crocodylus intermedius, respectively) show exceptionally high-pressure values (outside the 95% confidence interval), and Crocodylus johnsoni shows pressures expected of animals nearly a magnitude in size larger. Other ecomorphs show much lower and similar relative values. The arrow indicates the typical ultimate shear strength of bone. (B) Ancestral character-state reconstruction using squared-change parsimony of size-standardized caniniform pressures. Residual caniniform pressure values are color coded to MPa (squared-change parsimony; squared length = 19.491). Vertical scale is in relative time, with the outgroup/ingroup root arbitrarily set to 1.0. High relative pressures were achieved independently in Crocodylus intermedius, Gavialis gangeticus, and Crocodylus johnsoni. Uncolored branches represent taxa for which the caniniform teeth were shed or broken, and so pressure estimation was not possible.CC BY · PubMed Central
Figure 5 Linear regressions of residual caniniform tooth pressures, and residual molariform tooth pressures versus rostral proportion phylogenetic independent contrasts. The (A) residual caniniform, and (B) residual molariform regressions show the low correlation between these parameters after accounting for body mass and phylogenetic relatedness.
Figure 5 Linear regressions of residual caniniform tooth pressures, and residual molariform tooth pressures versus rostral proportion phylogenetic independent contrasts. The (A) residual caniniform, and (B) residual molariform regressions show the low correlation between these parameters after accounting for body mass and phylogenetic relatedness.CC BY · PubMed Central
Figure 6 Molariform pressure values for extant Crocodylia, their phylogenetic distribution, and inferred ancestral character states. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the relationship between body size and molariform pressure. Note that the range of values shows similar interspecific correspondence to the caniniform pressure data shown in Figure 4A. The arrow indicates the typical ultimate shear strength of bone. (B) Ancestral-state reconstruction using squared-change parsimony of size-standardized molariform pressures. Pressures are color coded to MPa. Vertical scale is in relative time, with the outgroup/ingroup root arbitrarily set to 1.0. The notable similarities between unrelated taxa and differences between related taxa illustrate the large amount of convergence for this trait among crocodilians.
Figure 6 Molariform pressure values for extant Crocodylia, their phylogenetic distribution, and inferred ancestral character states. (A) Members of the Alligatoridae are shown in blue, and members of the Crocodylidae+Gavialidae in green. The OLS regression equation describes the relationship between body size and molariform pressure. Note that the range of values shows similar interspecific correspondence to the caniniform pressure data shown in Figure 4A. The arrow indicates the typical ultimate shear strength of bone. (B) Ancestral-state reconstruction using squared-change parsimony of size-standardized molariform pressures. Pressures are color coded to MPa. Vertical scale is in relative time, with the outgroup/ingroup root arbitrarily set to 1.0. The notable similarities between unrelated taxa and differences between related taxa illustrate the large amount of convergence for this trait among crocodilians.CC BY · PubMed Central

Verification

Claims the paper makes

40 Own result · 35 Own interpretation · 17 No full text of the sources · 10 Own method · 10 No basis given · 8 Not checked yet · 2 Supported · 2 Sources not found · 1 Couldn’t confirm · accuracy not yet measured

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  • It may state two separate claims as one. 29
  • Probably not a substantial claim. 16
40 of 40
  • The paper relies on this without support.

    Extinct crocodilians followed the same body-mass-to-bite-force scaling relationship as extant species, so regression equations from extant taxa can predict fossil bite forces.

    “Bite forces can now be predicted in fossil crocodilians using the regression equations generated in this research.”
    “Crocodilian bite-force retention can be used to predict forces in other specimens and species, including taxa known only from fossils (see Materials and Methods).”
  • The paper relies on this without support.

    Crocodilians typically bite unilaterally, so the crowns of the prominent caniniform teeth initiate contact with prey during biting.

    “Because of their greater length relative to the adjacent teeth, and the propensity of crocodilians to bite unilaterally, the crowns of these teeth typically initiate contact with prey during biting.”
  • The paper relies on this without support.

    The caniniform and molariform teeth of crocodilians are composed of the same dental constituents, enamel and von Ebner's dentine.

    “We suspect the reason for the similarity is that both tooth types are composed of the same dental constituents (enamel and von Ebner's dentine) and must be able to damage, and yet sustain impacts with the same tissue types when feeding.”
  • The paper relies on this without support.

    The prominent molariform tooth position is where crocodilians primarily crush prey.

    “This is an ecologically relevant location since it is where these animals primarily crush prey.”
  • The paper relies on this without support.

    Crocodilians stereotypically seize prey contacted by the teeth and jaws as the head is swiped to the side.

    “Crocodilians stereotypically seize prey contacted by the teeth and jaws as the head is swiped to the side.”
  • The paper relies on this without support.

    Crocodilians process food on one side of the jaw.

    “They also process food on one side of the jaw.”
  • The paper relies on this without support.

    Unilateral crushing of prey at the molariform teeth commonly occurs with the head out of water in all species.

    “Additionally, unilateral crushing of prey at the molariform teeth commonly occurs with the head out of water in all species.”
  • The paper relies on this without support.

    Seizing of prey using the caniniform teeth often occurs with the head out of water.

    “Similarly, the seizing of prey using the caniniform teeth often occurs with the head out of water.”
8 of 8

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Metadata

Authors
Erickson, G. M., Gignac, P. M., Steppan, S. J., Lappin, A. K., Vliet, K. A., Brueggen, J. D., Inouye, B. D., Kledzik, D., & Webb, G. J. W.
Affiliations
  • Florida State University (US)
  • California State Polytechnic University (US)
  • University of Florida (US)
  • Charles Darwin University (AU)
Corresponding author
Erickson, Gregory M.
Venue
PLoS ONE
Published
2012-03-14
Volume
7
Issue
3
Pages
e31781
ISSN
1932-6203 (ISSN-L)
OpenAlex
W2048410549
Cited by
218
References
52
Citations by year
2026: 13 · 2025: 10 · 2024: 16 · 2023: 8 · 2022: 16 · 2021: 27 · 2020: 16 · 2019: 28 · 2018: 10 · 2017: 17 · 2016: 14 · 2015: 15 · 2014: 13 · 2013: 11 · 2012: 4
FWCI
10.25
Type
journal article
Language
en
Open access
Locations
  • PLoS ONE (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • CDU eSpace Institutional Repository (Charles Darwin University) (submittedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Funders
  • National Science Foundation (0841447, 0959029, 0623791)
  • National Geographic Society
Topics
  • Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
  • Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
  • Health Sciences › Medicine › Anatomy › Comparative Animal Anatomy Studies
Keywords
crocodilians, bite force, feeding mechanics, jaw morphology, body size, feeding evolution, fossil crocodilians, phylogenetic analysis, evolutionary success
MeSH terms
Alligators and Crocodiles, Animals, Biomechanical Phenomena, Bite Force, Body Weight, Biological Evolution, Jaw, Molar, Phylogeny, Predatory Behavior, Pressure, Skull, Tooth, Linear Models, Extinction, Biological, Ecological and Environmental Phenomena

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