Distinct evolutionary signatures underlie body shape diversity across deep-sea habitats
Data files
Oct 15, 2025 version files 318.10 KB
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dryad_package_Oct_14_2025.zip
312.33 KB
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README.md
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Abstract
The deep sea is known for extreme biological conditions such as high pressure, little to no solar light, and cold temperatures. Despite these challenges, deep-sea fishes have been shown to have higher body shape diversity than shallow-sea relatives. An open question is whether different habitats within the deep sea differentially contribute to this surprising phenotypic diversity. Here, we explore the joint effects of two major environmental dimensions in marine ecosystems, the benthic-pelagic axis and ocean depth, on phenotypic diversification in teleost fishes. Using measurements of body shape across nearly 3,000 species, we found that increasing ocean depth generally shifted axes of phenotypic evolution and promoted diversification for benthic, demersal, and pelagic fishes alike. However, body shape diversity and rates of body shape evolution did not scale evenly across these habitat divisions. For benthic fishes, the rate increased more strongly than diversity with ocean depth, while the reverse was true for pelagic fishes. Analyses of historical transition rates showed that routes to colonizing deep-pelagic habitats have been more variable than those for colonizing deep-benthic habitats, suggesting that independent invasions from different sources may help explain the diversity of deep-pelagic fishes without invoking high evolutionary rates. Relaxed selection on body shape may also explain this diversity, as suggested by the extreme range of forms found in the deep pelagic, coinciding with an elongation axis shared by all deep-sea fishes. Overall, our results reveal a mosaic of pathways through which body plan diversity has accumulated across a large vertebrate radiation, underscoring the importance of considering finer-scale habitat variation in broad-scale evolutionary studies.
Dryad DOI: https://doi.org/10.5061/dryad.z612jm6r1
Main file: dryad_package_Oct_14_2025.zip
Dryad package prepared by Elizabeth Santos (neé Miller) on October 14, 2025
Contact with questions: lizmiller2633@gmail.com
R scripts are designed to work with the R project "comp_methods.Rproj." Open this file first!
This R project keeps all folders within a common directory so the scripts can all "talk" to each other
Description of files:
Folder "data"
contains input files needed to run all analyses
"pruned_rab.tre" is a newick-formatted time-calibrated phylogenetic tree originally published by Rabosky et al. 2018, with taxa pruned to match sampling of marine species with body shape data (see Table S2).
"size_corrected_morph_withcombined_hab_cats.csv" contains body shape variables and habitat data that are input into all analyses.
- column B "tip_name" matches the tip label in the Rabosky et al. phylogeny ("pruned_rab.tre")
- Columns C–J contain eight body shape variables published by Price et al. 2022 (for methods of collection see Price et al. 2019). These variables have been size-adjusted using log-shape ratios (see Methods section)
- Column K "final_combined_habitat" merges the depth classification with the water-column-use classification (see below) with an underscore in the middle. There are nine categories in total after merging. (referred to as "nine-state scheme" throughout our manuscript)
- Column L "depth_cat" contains the categorical depth classifications for each marine species published by Martinez et al. 2021, with no modifications
- column M "water_col" contains the categorical water-column-use classifications for each marine species published by Friedman et al. 2020, with no modifications
- Column N "merged_benthic_dem_cat" is a reclassification of column K after subsuming the "benthic" category into the "demersal" category (here called simply 'demersal'). This is referred to as the 'six-state scheme' throughout our manuscript.
Folder "morphospace":
"plot morphospaces.R" is an R script that reproduces morphospace panels shown in Fig. 1 and Fig. S1
The folders "density_panels" and "density_panels_merged_demersal" are empty folders meant to corral outputs exported by this R script
Folder "phyloMANOVA":
"phyloMANOVA.R" is an R script that reproduces phylogenetic MANOVA analyses and exports results to this folder (see Tables S7, S8)
Folder "simmap":
"simmap.R" is an R script that reproduces SIMMAP analyses and exports results to this folder
Folder "hypervolumes":
"hypervolumes.R" is an R script that calculates observed hypervolume size and pairwise comparisons of hypervolume overlap among habitats (e.g., Fig. 3, Fig. S8, and red bars of Figs. S6 and S7)
The same script calculates distributions of permuted hypervolumes for comparison with the observed values (see Table S13 and Figs. S6–S7)
This script is modified from those published by Corn et al. 2022 (link to Zenodo repository within script)
Folder "geomorph_rates_disparity_comparison":
"geomorph disparity and rates.R" calculates phenotypic disparity and evolutionary rates by habitat (see Fig. 3, Tables S9–12, Fig. S8)
Folder "pairwise_angles":
The scripts "functions.R" and "run.R" (written by Sarah Friedman) are meant to work together to replicate the analyses comparing angles of primary axes of morphological variation (see Fig. 2, Figs. S3 and S4, Table S5 and S6)
The "functions.R" script contains custom functions setting up the analyses, and is read by the "run.R" script to control running the analyses
"output" folder is empty and meant to corral outputs associated with these scripts only
Scripts are modified from those used in Friedman et al. 2022 Syst. Biol.
Relevant references below:
Corn, K. A., S. T. Friedman, E. D. Burress, C. M. Martinez, O. Larouche, S. A. Price, and P. C. Wainwright. 2022. The rise of biting during the Cenozoic fueled reef fish body shape diversification. Proc. Natl. Acad. Sci. U.S.A. 119:e2119828119.
Friedman, S. T., S. A. Price, K. A. Corn, O. Larouche, C. M. Martinez, and P. C. Wainwright. 2020. Body shape diversification along the benthic–pelagic axis in marine fishes. Proceedings of the Royal Society B: Biological Sciences 287:20201053.
Friedman, S. T., M. L. Collyer, S. A. Price, and P. C. Wainwright. 2022. Divergent Processes Drive Parallel Evolution in Marine and Freshwater Fishes. Systematic Biology 71:1319–1330.
Martinez, C. M., S. T. Friedman, K. A. Corn, O. Larouche, S. A. Price, and P. C. Wainwright. 2021. The deep sea is a hot spot of fish body shape evolution. Ecology Letters 24:1788–1799.
Price, S. A., S. T. Friedman, K. A. Corn, C. M. Martinez, O. Larouche, and P. C. Wainwright. 2019. Building a Body Shape Morphospace of Teleostean Fishes. Integrative and Comparative Biology 59:716–730.
Price, S. A., S. T. Friedman, K. A. Corn, O. Larouche, K. Brockelsby, A. J. Lee, M. Nagaraj, N. G. Bertrand, M. Danao, M. C. Coyne, J. R. Estrada, R. Friedman, E. Hoeft, M. Iwan, D. Gross, J. H. Kao, B. Landry, M. J. Linares, C. McGlinn, J. A. Nguyen, A. G. Proffitt, S. Rodriguez, M. R. Rupp, E. Y. Shen, V. Susman, A. J. Tovar, L. L. J. Vary, K. L. Zapfe, and P. C. Wainwright. 2022. FishShapes v1: Functionally relevant measurements of teleost shape and size on three dimensions. Ecology 103:e3829.
Rabosky, D. L., J. Chang, P. O. Title, P. F. Cowman, L. Sallan, M. Friedman, K. Kaschner, C. Garilao, T. J. Near, M. Coll, and M. E. Alfaro. 2018. An inverse latitudinal gradient in speciation rate for marine fishes. Nature 559:392–395.
