Integrating biotic interactions in niche analyses unravels patterns of community composition in clownfishes
Data files
Apr 13, 2023 version files 46 MB
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3U_estimates_simplified.csv
23.73 KB
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amph_behavior.csv
990 B
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environmental_dataset.csv
18.15 MB
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GG_spatGLMmodel_predictions.csv
45.08 KB
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GS_spatGLMmodel_predictions.csv
47.52 KB
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interaction_matrix.csv
1.45 KB
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marine_regions.csv
6.30 MB
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niche_overlaps.csv
44.08 KB
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overall_spatGLMmodel_predictions.csv
44.61 KB
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README.md
23.34 KB
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spatial_results.csv
21.27 MB
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SS_spatGLMmodel_predictions.csv
47.15 KB
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summary_spatial.csv
3 KB
Sep 24, 2024 version files 46 MB
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3U_estimates_simplified.csv
23.73 KB
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amph_behavior.csv
990 B
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environmental_dataset.csv
18.15 MB
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GG_spatGLMmodel_predictions.csv
45.08 KB
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GS_spatGLMmodel_predictions.csv
47.52 KB
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interaction_matrix.csv
1.45 KB
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marine_regions.csv
6.30 MB
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niche_overlaps.csv
44.08 KB
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overall_spatGLMmodel_predictions.csv
44.61 KB
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README.md
21.48 KB
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spatial_results.csv
21.27 MB
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SS_spatGLMmodel_predictions.csv
47.15 KB
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summary_spatial.csv
3 KB
Sep 27, 2025 version files 880.93 MB
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data.zip
5.75 MB
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Rdata.zip
873.08 MB
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README.md
19.25 KB
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results.zip
2.08 MB
Sep 30, 2025 version files 880.93 MB
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data.zip
5.75 MB
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Rdata.zip
873.08 MB
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README.md
19.25 KB
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results.zip
2.08 MB
Mar 11, 2026 version files 1.41 GB
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data.zip
5.75 MB
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figures.zip
36.21 MB
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Rdata.zip
990.44 MB
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README.md
10.14 KB
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results.zip
377.78 MB
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tables.zip
4.41 KB
Abstract
Mutualistic interactions play a fundamental role in shaping species distributions, driving niche differentiation, and structuring communities. Yet their influence on realized niches and patterns of coexistence remains poorly understood. In clownfishes, mutualism with sea anemones underpins their biogeography and ecological success, with species classified as generalists or specialists according to their host specificity. However, the extent to which host availability constrains or expands clownfish niches has remained unclear. Here we used ecological niche models (ENMs) across the Indo-Pacific to examine how host sea anemones shape clownfish realized niches and community composition. We integrated occurrence data with climatic and habitat predictors and refined estimates by incorporating host niches, quantifying changes in niche breadth, position, and overlap. We further developed a multilayered framework to account for host use within niche quantification, allowing us to assess patterns of resource overlap in communities spanning generalist–specialist dynamics. Our results reveal that mutualistic associations strongly shape clownfish realized niches, with specialists experiencing greater niche constraints than generalists due to their reliance on a limited set of hosts. Host availability emerges as a key driver of community structure, producing high ecological niche overlap among species that is modulated by host partitioning. This host-mediated differentiation facilitates coexistence by reducing resource overlap, with generalist–specialist contrasts playing a central role in sustaining clownfish biodiversity. Our study highlights how biotic interactions mediate species ecological niches and shape community assembly in an iconic mutualistic system. Beyond clownfishes, our framework offers a transferable approach to incorporate interacting species into ENMs, improving ecological interpretations and informing conservation efforts in biodiversity hotspots and other mutualistic systems.
Data for: Integrating biotic interactions in niche analyses unravels patterns underlying community composition in clownfishes
1. General Information
Dataset Title
Data for the article:
Integrating biotic interactions in niche analyses unravels patterns underlying community composition in clownfishes
Dataset DOI
https://doi.org/10.5061/dryad.2bvq83bv8
Authors
Alberto Garcia Jimenez
Olivier Broennimann
Antoine Guisan
Théo Gaboriau
Nicolas Salamin
Author ORCID IDs
Alberto Garcia Jimenez – https://orcid.org/0000-0002-1532-8784
Olivier Broennimann – https://orcid.org/0000-0001-9913-3695
Antoine Guisan – https://orcid.org/0000-0002-3998-4815
Théo Gaboriau – https://orcid.org/0000-0001-7530-2204
Nicolas Salamin – https://orcid.org/0000-0002-3963-4954
Contact
Alberto Garcia Jimenez
Department of Computational Biology
University of Lausanne
Email:
agarcia26286@gmail.com
alberto.garciajimenez@unil.ch
2. Dataset Description
This dataset contains the data, scripts, model outputs, and figures used in the study:
Garcia Jimenez et al. – Integrating biotic interactions in niche analyses unravels patterns underlying community composition in clownfishes.
The study investigates how biotic interactions between clownfish species (Amphiprion spp.) and their host sea anemones influence ecological niche structure and spatial community composition across the Indo-Pacific.
The dataset includes:
- species occurrence datasets
- environmental predictors used in ecological niche modelling
- interaction matrices between clownfish and sea anemones
- niche overlap and spatial analysis results
- environmental niche models (ENMs)
- mutualism-refined niche models
- statistical model outputs
- reproducible analysis scripts
- figures and supplementary tables used in the manuscript
3. Repository Structure
The dataset is organized into compressed archives.
Dataset
├── data.zip
├── results.zip
├── Rdata.zip
├── figures.zip
├── tables.zip
└── scripts.zip
4. Recommended Software
The following free or open-source software can be used to view the files:
.csv — R, Python, LibreOffice Calc
.R — RStudio or R
.RDS — R
.pdf — any PDF reader
.tex — LaTeX editors such as TeX Live or Overleaf
.tif — QGIS, ArcGIS, or R packages such as terra
.shp/.dbf/.shx/.prj — QGIS, ArcGIS, or R package sf |Note: Shapefiles can be opened and used in any GIS software and in R or Python. A shapefile consists of multiple file types beyond the .shp (specifically, .cpg, .dbf, .prj, .sbn, and .sbx). The user only interacts directly with the .shp file but the other files need to be in the same directory.
.zip — any archive utility
Analyses were developed using:
R ≥ 3.6.3
RStudio 2021.09.2
5. Data Sources
Occurrence records were compiled from:
- GBIF (Global Biodiversity Information Facility)
- Reef Life Survey (RLS)
- OBIS (Ocean Biogeographic Information System)
- Hexacoral database
Environmental predictors were obtained from:
- GMED (Global Marine Environment Datasets)
- Bio-Oracle
Marine regions follow the Marine Ecoregions of the World (MEOW) classification.
6. Folder Descriptions
6.1 data.zip
This folder contains all primary input datasets used in the analyses.
data/
- amph_occ_env_final_dataset.csv
- anem_occ_env_final_dataset.csv
- interaction_matrix.csv
- old_interaction_matrix.csv
- selected_environmental_variables.csv
- marine_regions.csv
- marine_regions.shp
- marine_regions.dbf
- marine_regions.shx
- marine_regions.prj
- meow_ecos_df.csv
- qc/
Occurrence datasets
amph_occ_env_final_dataset.csv — Geo‑referenced occurrence records for clownfish species including environmental predictors.
anem_occ_env_final_dataset.csv — Geo‑referenced occurrence records for sea anemone host species including environmental predictors.
Dataset format:
species: clownfish species name (character)
x: longitude (decimal degrees)
y: latitude (decimal degrees)
Interaction matrices
interaction_matrix.csv — Species-named binary matrix describing documented clownfish–anemone mutualistic interactions.
old_interaction_matrix.csv — Previous version of the interaction matrix retained for reproducibility of earlier analyses.
Environmental predictors
selected_environmental_variables.csv — Environmental predictor variables extracted for all grid cells in the study region (~0.083° resolution).
Environmental predictor variables extracted for all grid cells in the study region
at approximately 0.083° spatial resolution. These variables were used in the
environmental niche modelling analyses.
Each row corresponds to a geographic grid cell.
Variables:
x: Longitude of the grid cell centre. Units: decimal degrees.
y: Latitude of the grid cell centre. Units: decimal degrees.
bo2dissolve: Mean dissolved oxygen concentration at the bottom of the water column. Units: mmol m-3.
bphosphate: Mean phosphate concentration in seawater. Units: µmol L-1.
chlarange: Range (max–min) of chlorophyll-a concentration across the year. Units: mg m-3.
parmean: Mean photosynthetically active radiation. Units: Einstein m-2 day-1.
dissolved.oxygen.mean: Mean dissolved oxygen concentration along the water column. Units: mmol m-3.
nitrate.mean: Mean nitrate concentration. Units: µmol L-1.
temperature.range: Annual range of sea surface temperature. Units: °C.
sstmean: Mean sea surface temperature. Units: °C.
sstrange: Range of sea surface temperature across the year. Units: °C.
Marine regions
marine_regions.shp/.dbf/.shx/.prj — GIS shapefile representing marine provinces used in spatial analyses.
marine_regions.csv — Tabular version of marine region classifications.
Variables:
x: Longitude of the grid cell centre. Units: decimal degrees.
y: Latitude of the grid cell centre. Units: decimal degrees.
province: Marine province assigned to the grid cell based on the MEOW biogeographic classification.
realm: Marine realm assigned to the grid cell based on the MEOW biogeographic classification.
QC datasets
qc/ contains summary datasets used during data preparation to assess overlap among occurrence data sources.
6.2 results.zip
Contains datasets produced as outputs of the analyses.
These include:
- ROU (Relative Occupancy of the environmental niche) metrics
- niche overlap datasets
- spatial analysis datasets
- statistical model outputs
- environmental niche models
- mutualism‑refined niche models
- raster predictions
- shapefiles of predicted distributions
Key datasets include:
ROU_data.csv
ROU_data_envhost_predictors.csv
ROU_data_host_predictors.csv
ROU_data_oldA.csv
niche_overlaps.csv
niche_overlaps_corrected.csv
spatial_results.csv
spatial_results_corrected.csv
summary_spatial.csv
with additional folders:
├── amph_EBMs_newA
├── amph_EBMs_oldA
├── amph_ENMs
├── anem_ENMs
├── raster_maps
└── shapefiles
containing environmental niche models to be loaded using NINA function load_model(folder) and predicted species distributions provided as raster GeoTIFF files (.tif) and shapefiles.
6.3 Rdata.zip
Contains serialized R objects used in the analyses.
Rdata/
- GG_spatialGLMM.RDS
- GS_spatialGLMM.RDS
- SS_spatialGLMM.RDS
- all_spatialGLMM.RDS
- amphEBMs.RDS
- amphEBMs_oldA.RDS
- amphENMs.RDS
- amphENMs_envhostpredictors.RDS
- amphENMs_glob.RDS
- anemENMs.RDS
- anemENMs_glob.RDS
These files contain fitted model objects and can be loaded in R using:
readRDS("filename.RDS")
6.4 scripts.zip
Contains all R scripts required to reproduce the analyses.
Scripts include:
0a_occ_datapreparation.R
0b_env_datapreparation.R
0c_marine_regions.R
0d.env_var_selection.R
1_reg_models.R
2_ROU_analysis.R
3_nicheOverlaps.R
4_spatialCompetition.R
4_spatialCompetition_corrected.R
GLM_and_moranItests.R
Models_evaluation.R
spatialGLMM.R
functions.R
Additional scripts generate figures and tables used in the manuscript.
6.5 figures.zip
Contains all main and supplementary figures used in the manuscript.
figures/
- Fig1–Fig5 (main manuscript figures)
- FigS1–FigS10 (supplementary figures)
- maps/ (species distribution maps derived from niche modelling analyses)
6.6 tables.zip
Contains supplementary tables in LaTeX format.
tables/
- TableS1_Cramers_test.tex
- TableS2_ROU_GvsS_summary.tex
- TableS3.Spatial GLMMs.tex
7. Missing Data Codes
NA — missing value
8. License
All files in this dataset are distributed under the CC0 1.0 Universal Public Domain Dedication, as required by Dryad.
9. Confirmation Regarding Licensing and Third‑Party Data
The authors confirm that all materials included in this Dryad submission are compatible with the CC0 license waiver required by Dryad.
Occurrence data used in the analyses were obtained from publicly accessible biodiversity databases (GBIF, OBIS, Reef Life Survey) and were processed and aggregated by the authors for analytical purposes.
Environmental predictors were derived from publicly available environmental datasets (GMED and Bio‑Oracle).
The versions included in this repository consist of derived analytical datasets, model outputs, and figures generated by the authors.
All figures included in figures.zip were produced by the authors using the scripts included in this dataset. The authors confirm that these figures may be distributed under the CC0 waiver applied by Dryad.
The authors confirm that the dataset does not contain copyrighted material redistributed in its original form and that all files included here are compatible with the CC0 public domain dedication required for Dryad deposits.
Occurrences, environmental variables and all input data for the analyses have been collected from online databases and open-source datasets.
All analyses are done in R. Developed functions are provided in form of a package called NINA.
Changes after Apr 13, 2023:
Changes after Sep 24, 2024:
Changes after Sep 27, 2025:
Changes after Sep 30, 2025:
Changes after March 1, 2026:
These additions and updates were implemented following the peer-review
process of the associated manuscript.
The revisions mainly aim to:
(1) allow comparison between analyses based on the previous interaction
matrix (“old A”) and the updated interaction matrix,
(2) improve transparency and reproducibility by including the analysis
scripts directly in the Dryad repository,
(3) provide additional supporting analyses requested by reviewers,
and (4) include the final figures and supplementary tables used in the manuscript.
NEW FILES AND FOLDERS
data/
- qc/
Quality-control datasets summarizing overlap among occurrence records
from different biodiversity databases used during data preparation.
- old_interaction_matrix.csv
Previous version of the clownfish–anemone interaction matrix retained
to allow comparison between analyses using the original and the updated
interaction matrices.
results/
- ROU_data_oldA.csv
Relative Occupancy of the environmental niche (ROU) metrics calculated
using the previous interaction matrix ("old A"), allowing comparison
with the updated analyses reported in the manuscript.
- anem_regtable.csv
Table summarizing anemone distribution ranges used to validate that the
predicted environmental niche model distributions are consistent with
current literature descriptions.
- amph_ENMs/
Folder containing environmental niche models for clownfish species
generated using the NINA modelling framework.
figures/
Main manuscript figures (Fig1–Fig5) and supplementary figures
(FigS1–FigS10) generated during the final stage of the revision process.
- figures/maps/
Species distribution maps derived from mutualism-refined environmental
niche models.
tables/
- TableS1_Cramers_test.tex
- TableS2_ROU_GvsS_summary.tex
- TableS3_Spatial_GLMMs.tex
Supplementary tables in LaTeX format used in the manuscript appendix.
scripts/
All analysis scripts required to reproduce the analyses.
These scripts were previously available only through the project GitHub
repository and are now included directly in the Dryad dataset to ensure
long-term archival reproducibility.
Rdata/
- amphEBMs_oldA.RDS
Serialized R object containing environmental niche models fitted using
the previous interaction matrix (“old A”), enabling direct comparison
with the updated models.
UPDATED FILES
Several results datasets and figures were updated following reviewer
comments and the incorporation of the alternative interaction matrix
analysis.
These updates affect:
results/
- ROU_data_corrected.csv
- niche_overlaps_corrected.csv
- spatial_results_corrected.csv
scripts/
Updated scripts generating the revised analyses and figures.
- Garcia Jimenez, Alberto et al. (2023), Implementation of biotic interactions in niche analyses unravels the patterns underneath community composition in clownfishes, , Article, https://doi.org/10.5281/zenodo.7668897
- Jimenez, Alberto Garcia; Broennimann, Olivier; Guisan, Antoine et al. (2023). Implementation of biotic interactions in niche analyses unravels the patterns underneath community composition in clownfishes [Preprint]. Authorea, Inc.. https://doi.org/10.22541/au.167725089.96218529/v1
- Jiménez, Alberto García; Guisan, Antoine; Broennimann, Olivier et al. (2023). Integrating Biotic Interactions In Niche Analyses Unravels Patterns Of Community Composition in Clownfishes [Preprint]. Cold Spring Harbor Laboratory. https://doi.org/10.1101/2023.03.30.534900
- Garcia Jimenez, Alberto; Broennimann, Olivier; Guisan, Antoine et al. (2025). Integrating biotic interactions in niche analyses unravels patterns of community composition in clownfishes. Zenodo. https://doi.org/10.5281/zenodo.7668896
- Garcia Jimenez, Alberto; Broennimann, Olivier; Guisan, Antoine et al. (2025). Integrating biotic interactions in niche analyses unravels patterns of community composition in clownfishes. Zenodo. https://doi.org/10.5281/zenodo.17226218
