Phylogeny, systematics and evolution of mimicry patterns in Neotropical limenitidine butterflies
Data files
Sep 04, 2025 version files 61.39 MB
-
Paez_et_al_SYEN_mimpatt_evol_datasets_R1.zip
25.51 KB
-
Paez_et_al_SYEN_PhylogeneticAnalyses_R1.zip
61.35 MB
-
README.md
10.70 KB
Abstract
The Neotropical butterfly genus Adelpha Hübner exhibits remarkable species diversity and striking convergence in wing colour patterns, potentially explained by mimicry, making it an exceptional model for exploring trait evolution and its relationship with speciation. To date, unresolved phylogenetic relationships hinder a comprehensive understanding of the evolutionary biology of the genus. Using a novel multi-marker dataset combining one mitochondrial and 15 nuclear gene fragments, we generate the most comprehensive phylogeny of the genus Adelpha to revisit its systematics and investigate the evolution of mimicry colour patterns. Our data set encompasses 83 of the 87 known extant species and six Limenitis species that were recently excluded from Adelpha (134 of c. 160 subspecies in total), collectively displaying 14 distinct mimicry patterns. We provide conclusive evidence that corroborates previous work on the polyphyly of Adelpha as historically conceived, and describe the genus Adelphina Páez & Willmott n. gen. to stabilize the nomenclature, both genera representing Neotropical limenitidines. The comprehensive phylogeny provided in this study lays a solid foundation for future research into the processes driving diversification within these species interacting through mimicry. Ancestral character state reconstruction reveals the gradual evolution of the mimicry pattern. The more common mimicry pattern IPHICLUS (forewing with orange subapical spot and white band) is inferred as ancestral, but repeated convergent evolution is also recovered. Evolutionary convergence is also observed for the second most abundant mimicry pattern, COCALA (orange-white banded) mimicry patterns. Increased rates of mimicry pattern evolution are also found toward the equator. These results underscore the complexity of mimicry evolution in the Neotropical limenitidines, i.e., Adelpha and Adelphina, emphasizing the need to explore its interplay with other biotic and abiotic factors.
Dataset DOI: 10.5061/dryad.7h44j104b
Description of the data and file structure
This README file was generated on 14-03-2025 by Erika Paez Vargas
This dataset, integral to our research paper, contains crucial data for understanding our key findings. It includes phylogenetic analysis data and results, raw data used to assess mimicry pattern evolution along with their R scripts, as well as supplementary materials and appendix data.
Files and variables
Below is a summary of the included files:
1. Phylogenetic and dating analyses
Folder: Paez_et_al_SYEN_PhylogeneticAnalyses_R1.zip
This folder contains the dataset and results of the topological tests, molecular phylogenetic and dating analyses.
- AU_test folder: the files and results to perform topological tests and ML analyses under IQ-TREE (v3.0.1).
- BEAST folder: the files to perform molecular dating analyses (using Bayesian relaxed clocks) using BEAST software package (v1.10.4), and results.
- IQTREE - gene tree analyses folder: files to perform molecular phylogenetic analyses per gene under IQTREE ( IQ-TREE v3.0.1), and tree file results for 1 mitochondrial gene and 15 nuclear genes.
- IQTREE2 - unconstrained folder: files to perform ML analyses under IQTREE (v2.2.2.7) with all subspecies included.
2. Reconstruction of the evolution of mimicry wing colour patterns
Folder: Paez_et_al_SYEN_mimpatt_evol_datasets_R1.zip
This folder contains the raw datasets and the tree file to perform all the analyses for the mimicry pattern evolution.
- Adelpha.pattern_list.txt: raw data for the ancestral state reconstruction analysis.
- Adelpha_Adelphina_Limenitis.txt: raw data for the ancestral state reconstruction analysis (Limenitidinae).
- Adelpha_node_states.txt: raw data for the ancestral state reconstruction analysis (Adelpha + Adelphina)
- lowland_Adelpha_node_states.txt: raw data for the ancestral state reconstruction analysis (only Adelpha)
- Adelpha_phylo.nwk: phylogenetic tree used in the ancestral state reconstruction analysis.
- Adelpha_tip_data.txt: raw data for the ancestral state reconstruction analysis.
- Adelpha_macroecol_traits.txt: raw data for the macroecological traits analyses using Phylogenetic Generalized Linear Models (PGLS).
- Adelpha_tip_data_Dstat.txt: raw data for the phylogenetic signal analysis (D-statistic).
- Adelpha_traits_plgs_2024: It summarizes ecological and evolutionary traits of various species in a dataset, with each row representing a different species (e.g., A.abia, A.abyla, etc.). This can be used to explore patterns in mimicry, niche breadth, geographic distribution, and evolutionary rates across species.
3. Electronic supplementary materials
Folder: Paez_et_al_SYEN_ESM_0309
- Figure S1: Estimation of ancestral character states for mimicry colour patterns in Adelpha + Adelphina + Limenitis showed that IPHICLUS colour pattern was most likely to be the state of the common ancestor of the Neotropical limenitidines Adelpha and Adelphina species.
- Figure S2: Estimation of ancestral character states for mimicry colour patterns only in Adelpha (lowland) species showed that IPHICLUS colour pattern was most likely to be the state of the common ancestor.
- Figure S3: A comparison of tree topologies from recent molecular phylogenetic studies, simplified to clades relevant to generic classification.
- Table S1: List of sequences including Voucher and GenBank accession numbers for the sixteen gene fragments.
- Table S2: Partition scheme inferred by the Modelfinder analysis of the specimen-level dataset.
- Table S3: Results of the AU-tests carried out with IQ-TREE.
- Table S4: Results for mimicry rate evolution analyses.
- Table S5: Summary of relevant limenitidine taxa included within recent molecular phylogenetic studies, a suggested generic-level classification, type species of genera, and inferred ages of genera from the present study.
4. Appendix files
Folder: Paez_et_al_SYEN__Appendix_R1.zip
This folder contains appendix files:
- Appendix S1: Details on generation of the transcriptomic data
- Appendix S2: Explanation for the calculation of the distribution maps for Adelpha species and macroecological traits.
- Appendix S3: Macroecological traits raw dataset.;
Code/software
All mimicry pattern evolution analyses were performed using R software.
-
Programs needed:
-
BayesTraits v4.0
Pagel, M., Meade, A., & Barker, D. (2004). Bayesian estimation of ancestral character states on phylogenies. Systematic Biology, 53(5), 673–684. https://doi.org/10.1080/10635150490522232
-
BEAST v1.10.4
Suchard, M. A., Lemey, P., Baele, G., Ayres, D. L., Drummond, A. J., & Rambaut, A. (2018). Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10. Virus Evolution, 4(1), vey016.
-
Codoncode Aligner (v3.7.1.1, CodonCode Corporation, http://www.codoncode.com/)
-
IQ-TREE v2.2.2.7
Minh, B. Q., Schmidt, H. A., Chernomor, O., Schrempf, D., Woodhams, M. D., von Haeseler, A., & Lanfear, R. (2020). IQ-TREE 2: New Models and Efficient Methods for Phylogenetic Inference in the Genomic Era. Molecular Biology and Evolution, 37(5), 1530–1534. https://doi.org/10.1093/molbev/msaa015
-
ModelFinder
Kalyaanamoorthy, S., Minh, B. Q., Wong, T. K. F., von Haeseler, A., & Jermiin, L. S. (2017). ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14(6), 587–589. https://doi.org/10.1038/nmeth.4285
-
Sequence Capture Processor (SECAPR) pipeline (http://htmlpreview.github.io/?https://github.com/AntonelliLab/seqcap_processor/blob/master/docs/documentation/main_doc.html)
-
Tracer v1.7.2
Rambaut, A., Drummond, A. J., Xie, D., Baele, G., & Suchard, M. A. (2018). Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67(5), 901–904. https://doi.org/10.1093/sysbio/syy032
-
-
Analysis Scripts:
Folder: Paez_et_al_SYEN_scripts_pattern_evolution_R1.zip
All anaylses were performed using R software.
- R-ASRPLOT_2025: Script for Ancestral State Reconstruction analyses of wing colour patterns
- R-D_stat_2025: Script for D statistics analyses
- R-PGLS_2025: Script for PGLS. wing colour pattern rates of evolution correlated to macroecological traits.
- R-TipRatesEvol_2025: Script for wing colour pattern evolution analyses
-
Loaded packages for R scripts:
-
ade4
Thioulouse J, Dray S, Dufour A, Siberchicot A, Jombart T, Pavoine S (2018). Multivariate Analysis of Ecological Data with ade4. Springer. doi:10.1007/978-1-4939-8850-1
-
ape
Paradis E, Schliep K. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics. 2019;35(3):526–8.
-
caper
Orme, D., Freckleton, R.P., Thomas, G.H., Petzoldt, T., Fritz, S. & Isaac, N. (2013) Caper: Comparative analyses of phylogenetics and evolution in R. R package version 0.5.2
-
dplyr
Wickham H, François R, Henry L, Müller K, Vaughan D (2023). dplyr: A Grammar of Data Manipulation. R package version 1.1.4, https://CRAN.R-project.org/package=dplyr.
-
geiger
Pennell, M. W., J. M. Eastman, G. J. Slater, J. W. Brown, J. C. Uyeda, R. G. FitzJohn, M. E. Alfaro, and L. J. Harmon {2014} geiger v2.0: an expanded suite of methods for fitting macroevolutionary models to phylogenetic trees. Bioinformatics, 30, 2216-2218
-
ggimage
Yu G (2023). _ggimage: Use Image in 'ggplot2'_. R package version 0.3.
-
ggplot2
H. Wickham. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York, 2016.
-
ggtree
Guangchuang Yu. Using ggtree to visualize data on tree-like structures. Current Protocols in Bioinformatics, 2020, 69:e96. doi: 10.1002/cpbi.96
-
ggtreeExtra
S Xu, Z Dai, P Guo, X Fu, S Liu, L Zhou, W Tang, T Feng, M Chen, L Zhan, T Wu, E Hu, Y Jiang, X Bo, G Yu. ggtreeExtra: Compact visualization of richly annotated phylogenetic data. Molecular Biology and Evolution 2021, 38(9):4039-4042. doi: 10.1093/molbev/msab166
-
MASS
Venables, W. N. & Ripley, B. D. (2002) Modern Applied Statistics with S. Fourth Edition. Springer, New York. ISBN 0-387-95457-0
-
mvtnorm
Genz A, Bretz F (2009). Computation of Multivariate Normal and t Probabilities, series. Lecture Notes in Statistics. Springer-Verlag, Heidelberg. ISBN 978-3-642-01688-2.
-
phangorn
Schliep K.P. 2011. phangorn: phylogenetic analysis in R. Bioinformatics, 27(4) 592-593
-
phytools
Revell, L. J. (2024) phytools 2.0: an updated R ecosystem for phylogenetic comparative methods (and other things). PeerJ, 12, e16505.
-
picante
S.W. Kembel, P.D. Cowan, M.R. Helmus, W.K. Cornwell, H. Morlon, D.D. Ackerly, S.P. Blomberg, and C.O. Webb. 2010. Picante: R tools for integrating phylogenies and ecology. Bioinformatics 26:1463-1464
-
RColorBrewer
Neuwirth E (2022). RColorBrewer: ColorBrewerpalettes. R package version 1.1-3https://CRAN.R-project.org/package=RColorBrewer
-
reshape2
Hadley Wickham (2007). Reshaping Data with the reshape Package. Journal of Statistical Software, 21(12), 1-20. URL http://www.jstatsoft.org/v21/i12/
-
scales
Wickham H, Pedersen T, Seidel D (2025). scales: Scale Functions for Visualization. Rpackage version 1.4.0, https://CRAN.R-project.org/package=scales.
-
tidyr
Wickham H, Vaughan D, Girlich M (2024). tidyr: Tidy Messy Data. package version 1.3.1https://CRAN.R-project.org/package=tidyr.
-
TreePar
Stadler T (2015). TreePar: Estimating Birth and Death Rates Based on Phylogenies. R package version 3.3
-
TreeSim
Stadler T (2019). TreeSim: Simulating Phylogenetic Trees. R package version 2.4. https://CRAN.R-project.org/package=TreeSim
-
Access information
Other publicly accessible locations of the data:
- Genbank
Data was derived from the following sources:
- NA
