Survey of tooth marked hadrosaurid (Dinosauria: Ornithischia) jugals from the Late Cretaceous of Alberta, implications for feeding behavior
Data files
Jun 03, 2026 version files 77.34 KB
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README.md
14.18 KB
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Supplementary_Code_and_Tables-_jugal_analysis.zip
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Abstract
Paleopathologies and tooth marks observed on bones have long been used for behavioral interpretations of extinct organisms, including non-avian dinosaurs. However, our understanding of this behavior is relatively limited by the rarity of large-scale surveys of recovered material. Here, we present a systematic survey of tooth marks across 148 hadrosaurid jugals from various Late Cretaceous formations of Alberta, Canada. Across 20 of the elements (13.5%), we observed 48 tooth marks, restricted to the lateral surface, 40% of which were located on the middle jugal body and showed no preferred orientation of the mark long axis. Across taphonomic modes, tooth marks are most common on isolated specimens, followed by bonebed specimens, with few articulated/semi-articulated/associated specimens showing marks. The proportion of tooth marked to non-tooth marked jugals is comparable to tooth marking frequencies observed for modern mammalian carnivores. While tooth marks were likely produced post-mortem during feeding, it is unclear whether tooth marks were produced when the jugal was removed to access underlying adductor musculature or during the removal of extraoral tissues or a combination of these behaviors. Future large-scale surveys of hadrosaurid cranial elements could help clarify which of these explanations is more likely and elucidate Cretaceous carnivore carcass processing behaviors. Our survey also identified an additional two specimens showing elongate healed marks extending anteroposteriorly across the lateral surface of the jugal body that are of unknown, but potentially the same, origin. Further specimens with similar paleopathologies could help decipher a likely mechanism of injury.
Dataset DOI: 10.5061/dryad.fqz612k6x
Description of the data and file structure
Supplementary tables are provided as individual tabs in a singular excel file which is then read into the R programming environment by the included R qmd file for the various analyses as described in the main text. The excel file containing the supplementary data and the R qmd file are provided together in a single zip file so that they can be easily saved to the same file folder which will prevent errors while reading in the data into the R environment.
Measurements provided in supplementary tables were taken from specimen photos using ImageJ /silicone peels using a ruler. Measurements, categories and modifiers were taken following Wyenberg-Henzler et al. (2024). Primary, secondary and tertiary categories and curvature modifiers are defined according to the Category-Modifier (CM) system of Wyenberg-Henzler et al. (2024). External and internal textural modifiers applied to tooth marks are defined under the CM system. For paleopathological marks and possible feeding-associated damage to the postorbital process, external and internal textural modifiers are simply recorded as "healing" or "damage" respectively.
Wyenberg-Henzler, T.C.A., Fowler, D.W., Currie, P.J., and Sullivan, C., 2024, The category-modifier system: a hierarchical classification scheme for vertebrate tooth marks: Paleobiology, v. 50, p. 1–19.
Institutional abbreviations used in the supplementary data and figures are as follows:
UALVP - University of Alberta Laboratory of Vertebrate Palaeontology, Edmonton, Alberta, Canada
TMP - Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada
Files and variables
File: Supplementary_Code_and_Tables-_jugal_analysis.zip
Description: Zip file containing an excel sheet with supplementary data tables (in tabs) containing specimen information, raw measurements, and statistical results for the tooth marks considered in this study and corresponding R code qmd file. Note: excel data file and R code qmd file should be saved in the same folder in order to prevent errors while reading the data into R.
Supplementary table 1 contains survey information on the taxonomic identity, locality information, relative completeness and whether or not jugal elements examined preserve tooth marks, paleopathologic marks and/or associated damage.
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specimen – specimen number for the original element at its corresponding institution
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taxa, genus – taxonomic identity of element
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element/element specifics - information regarding type of element
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state.of.element – relative completeness of the element; can be fragment, partial or mostly complete/complete as described in the main text
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body.region, element – element identification
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side – what side of the body the element is from; left or right; question mark denotes uncertainty in side identification
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formation – geologic formation the element was recovered from if applicable
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preservation.mode – context from which the jugal was collected; can be unknown, bonebed, isolated or associated/ articulated/ semi-articulated; question marks before a term indicates uncertainty
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bonebed – name of the bonebed (if applicable) the specimen was collected from
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locality – name of general locality specimen was collected from (for TMP, this is taken from the online database)
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pathologic.or.tooth.marked – if a jugal has one or more tooth mark(s)/paleopathologic mark(s); recorded as a yes or no
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tooth.marked – if a jugal has one or more tooth mark(s); recorded as a yes or no
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types.of.features – types of modifications a jugal preserves; either "none", one of three possible categories, "tooth marks" (if tooth-marked only), "damage" (if only damage to postorbital process), "pathologic" (if paleopathologic only), or a combination of any of these three categories
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number of tooth marks – number of tooth marks/paleopathologic marks observed on a given jugal
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notes – any additional notes
Supplementary table 2 contains a list of the silicone peels taken for this study, the location the silicone peels were taken from, numbers for the original specimens that the peels were taken from and corresponding specimen numbers for those peels.
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specimen – specimen number for the original element from which the silicone peel was taken from
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silicone peel – institutional number assigned for silicone peel taken from original specimen
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formation – geologic formation the element was recovered from if applicable
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taxa, genus – taxonomic identity of element
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preservation.mode – context from which the jugal was collected; can be unknown, bonebed, isolated or associated/ articulated/ semi-articulated; question marks before a term indicates uncertainty
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bonebed – name of the bonebed (if applicable) the specimen was collected from
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state.of.element – relative completeness of the element; can be fragment, partial or mostly complete/complete as described in the main text
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body.region, element, element specifics – element identification
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element/element specifics - information regarding type of element
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side – what side of the body the element is from; left or right; question mark denotes uncertainty in side identification
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surface – the surface of the element the silicone peel was taken from
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placement – the general location on the element surface that the silicone peel was taken from
Supplementary table 3 contains measurements and modifiers. specimen – specimen number for the original element at its corresponding institution
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silicone peel – institutional number assigned for silicone peel taken from original specimen
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formation – geologic formation the element was recovered from if applicable
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taxa, genus – taxonomic identity of element
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element/element specifics - information regarding type of element
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preservation.mode – context from which the jugal was collected; can be unknown, bonebed, isolated or associated/ articulated/ semi-articulated; question marks before a term indicates uncertainty
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state.of.element – relative completeness of the element; can be fragment, partial or mostly complete/complete as described in the main text
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body.region, element, element specifics – element with tooth marks/paleopathologic marks
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side – what side of the body the mark is observed on; left or right; question mark denotes uncertainty in side identification
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surface – the surface of the element the tooth mark(s)/paleopathologic mark(s) was/were identified on
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angle measuring surface – anatomical view of the element from which the orientation of tooth mark(s)/paleopathologic mark(s) was/were taken
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placement, specific.placement – the location of the tooth mark(s)/paleopathologic mark(s) on the element if noted in original publication
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label – number used to identify individual tooth marks in figures in the main text and in supplementary figures 1-2.
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length – mark length in mm representing the greatest linear dimension (measured parallel the bone surface) of a given tooth mark
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width – mark width in mm representing the greatest value of the dimension of a given tooth mark parallel to the bone surface but perpendicular to the length
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depth – mark depth in mm representing the greatest value of the dimension perpendicular to the bone surface for a given tooth mark
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edge modifier, internal modifier – the external/internal textural modifier assigned to a given tooth mark under the Category-Modifier system
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bone layer modified – the deepest bone layer modified by a given tooth mark
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number of curves – number of curves identified for a given elongate tooth mark
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c1, c2, c3, c4 – curvature measurements measured for a given tooth mark. Curvature is the angle defined by three points situated along one edge: the two ends of the edge, and the point of greatest curvature along the edge’s length. When a mark is approximately uniformly curved along its length, the third point is placed at the midpoint of the mark. For marks comprising two or more curves with different orientations, the transition points between successive curves are determined by visual estimation
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primary.category, secondary.category, tertiary.category – the primary/secondary/tertiary category assigned to a given tooth mark under the Category-Modifier system
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curve.category – the curvature modifier assigned to a given mark under the Category-Modifier system based on measured curvature
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angle (jugal anteroposterior axis) – angle between the line segment that defines the length, and the anteroposterior axis of the jugal, standardized to the left lateral surface
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standardized.jugal.axis – description of the original axis taken for the element that was used to reference the orientation of tooth mark measurements
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angle (skull anteroposterior axis) – angle used to rotate isolated jugals to match the orientation of a jugal on an articulated hadrosaurid skull in left lateral view
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tooth mark identification confidence – relative certainty with which a given mark represents a tooth mark/paleopathologic mark or associated damage
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notes – any additional notes pertaining to measurements or specimen
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angle between jugal anteroposterior axis and skull anteroposterior axis (jugal axis is clockwise of skull axis) – angle used to rotate isolated jugals to match the orientation of a jugal on an articulated hadrosaurid skull in left lateral view
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add angle difference – raw, re-calculated angle obtained for marks on jugals where the jugal is now rotated to its articulated position
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check angle is positive, if not, find positive angle by adding 180 – re-calculated angle obtained for marks on jugals where the jugal is now rotated to its articulated position, standardized to a positive equivalent angle between 0 and 180
Category-Modifier system is described in Wyenberg-Henzler, T.C.A., Fowler, D.W., Currie, P.J., and Sullivan, C., 2024, The category-modifier system: a hierarchical classification scheme for vertebrate tooth marks: Paleobiology, v. 50, p. 1–19.
Supplementary tables 4-5 contains the results of the omnibus and follow-up pairwise permuted t-tests log-transformed mark dimensions across various categories. Significant p-values are in bold. "Nm=" denotes number of tooth marks in each category. These tables are exported and formatted from results provided by the R qmd code file.
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Sum sq - sum of squares, returned from omnibus test conducted in R..
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Df - degree of freedom, returned from omnibus test conducted in R.
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F value - statistical F-value returned from omnibus test conducted in R.
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p-value - the p-value exported from the function; for pairwise comparisons in Supplementary Table 5, the p-values have been corrected for multiple comparisons using the Holm correction method as described in the main text
SUMMARY tab contains a breakdown of tooth mark distributions across various categories (e.g., number of left vs right jugals with tooth marks) which are discussed in the main text.
package.citations tab contains a list of the key R packages, version numbers for those packages and corresponding citations for the packages used in data analyses run in the R qmd file.
R qmd file used in the data analyses presented in the main text. This file was created and ran using R v. 4.5.1.
File: Supplementary_figures-_jugal_individual_maps.pdf (Zenodo)
Description: pdf containing supplementary figures 1 and 2 which depict individual mark maps for each specimen preserving tooth marks/associated damage/paleopathological marks. The numbers beside each tooth mark correspond to the tooth mark labels provided in the excel file containing supplementary data tables. These mark maps were manually generated from photographs of the specimens in Inkscape v2.0.
File: Supplementary_text_jugal_paper.docx (Zenodo)
Description: Word file containing detailed discussion exploring possible mechanisms of injury for paleopathological marks described in the main text. This file also contains a discussion of the third possible paleopathological jugal (FMNH UC 1479).
Code/software
Software:
Inkscape v2.0
ImageJ
R v. 4.5.1 via RStudio - qmd file contains code and user-defined functions which construct rose diagrams and other plots displayed in the main text. This code also runs various statistical analyses as described in the main text.
R packages:
Posit team. 2025. RStudio: Integrated development environment for r. Boston, MA: Posit Software, PBC.
Wilke CO. 2025. cowplot: Streamlined plot theme and plot annotations for “ggplot2”.
Aphalo PJ. 2025. ggpp: Grammar extensions to “ggplot2”.
Kassambara A. 2025. ggpubr: “ggplot2” based publication ready plots.
Schauberger P, Walker A. 2025. openxlsx: Read, write and edit xlsx files.
Pedersen TL. 2025. patchwork: The composer of plots.
Wickham H. 2011. The split-apply-combine strategy for data analysis. Journal of Statistical Software 40:1–29.
Wickham H. 2007. Reshaping data with the reshape package. Journal of Statistical Software 21:1–20.
HERVE M. 2025. RVAideMemoire: Testing and plotting procedures for biostatistics.
Wickham H, Henry L, Pedersen TL, Luciani TJ, Decorde M, Lise V. 2025. svglite: An “SVG” graphics device.
Wickham H, Averick M, Bryan J, Chang W, McGowan LD, François R, Grolemund G, Hayes A, Henry L, Hester J, Kuhn M, Pedersen TL, Miller E, Bache SM, Müller K, Ooms J, Robinson D, Seidel DP, Spinu V, Takahashi K, Vaughan D, Wilke C, Woo K, Yutani H. 2019. Welcome to the tidyverse. Journal of Open Source Software 4:1686. DOI: 10.21105/joss.01686.
Dragulescu A, Arendt C. 2020. xlsx: Read, write, format excel 2007 and excel 97/2000/XP/2003 files.
