Data from: Inferring time-to-speciation from hybrid zone analysis informs assessments of taxonomic inflation
Data files
May 05, 2026 version files 180.61 MB
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Gippner_dataset.zip
180.55 MB
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README.md
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Abstract
Measuring gene flow across hybrid zones can provide a direct evaluation of reproductive isolation (RI) between evolutionary divergent lineages. Geographic-explicit modelling of gene flow across hybrid zones, known as cline analysis, thus offers a gold standard for species delimitation under the biological species criterion. We here relate divergence time to postzygotic RI as inferred from hybrid zone cline width from studies on amphibians, reptiles, birds, mammals, and insects. The data confirm an overall significant negative correlation between cline width and divergence time (within-clade significant only in amphibians). Out of 108 hybrid zones, hybridizing taxa were younger than 10 million years (Ma) in 104 cases, and younger than 7 Ma in 94 cases, implying that older lineage pairs are usually reproductively isolated. In amphibians, the correlation suggested that shallow and steep hybrid zones, characteristic of subspecies and species, are found with divergence times of <2.6 Ma and >7 Ma, respectively. In all taxa, we revealed a trend of such young lineages being increasingly described as new species over time by simulating past sets of taxonomically named species. Overall, the inferred proportion of species with post-Pliocene ages was highest in birds (27.3%) and mammals (21.8%), but lower in amphibians and squamates (<10%). In amphibians, we document a 56% increase from 1980 to 2010 in the number of newly described species that fall below the hybrid zone-derived divergence reference point of 2.6 Ma. While this trend so far impacts only a small proportion of the total amount of amphibian species, we commend that taxonomic hypotheses that delimit Pleistocene-aged lineages as species, or conversely, that consider lineages with Miocene ages or older as conspecific, should be carefully interpreted to avoid taxonomic inflation and deflation.
https://doi.org/10.5061/dryad.wwpzgmsv2
Description of the data and file structure
The dataset was created to relate divergence time to postzygotic RI as inferred from hybrid zone cline width from studies on amphibians, reptiles, birds, mammals, and insects. For this, literature data was screened to collect cline width data, divergence time estimates, and dispersal rates. For amphibians, squamates, mammals, birds, a subgroup of bony fish, lepidopterans, and mollusks, past sets of taxonomically named species were simulated, and their divergence time was calculated using the software TaxoInflationAnalyzer.
Data Package Structure
All files associated with this dataset are provided within a single compressed archive:
- Gippner_dataset.zip
After extraction, the directory structure is as follows:
- Hybrid_zone_data.xlsx
- TInflA/
- results/
- species_lists/
- timetrees/
- R/
File Access
- The compressed archive (.zip) must be extracted before use
- After extraction, all files can be accessed using standard software:
- .xlsx files: Microsoft Excel or compatible programs
- .txt, .tsv, .tre files: any standard text editor
- .R files: R or RStudio
All analyses can be reproduced after extracting the archive and running the R script with the provided input files.
Files and variables
Overview
The dataset consists of:
- A central Excel file: Hybrid_zone_data.xlsx
- A results/ directory containing processed outputs (e.g., ####_out.txt)
- An R/ directory containing scripts and derived data used for analyses
File: Hybrid_zone_data.xlsx
Sheet: All_hybrid_zones
Description: Master list of all hybrid zones included in the dataset.
Variables:
- HZ_ID: Unique identifier for each hybrid zone
- organism: Broad organism category (e.g., bird, insect)
- Taxon_1: First taxon in the hybrid zone
- Taxon_2: Second taxon in the hybrid zone
- reference: Source publication describing the hybrid zone
Sheet: All_clines
Description: Summary of clines measured across hybrid zones and transects.
Variables:
- HZ_ID: Hybrid zone identifier
- Transect_ID: Identifier for sampling transect
- Hybridizing_taxa: Taxa involved
- Group: Taxonomic group
- Transect_Lit_ID: Transect-Literature reference ID
- Number_marker: Number of genetic markers used
- Number_marker_cat: Categorical bin of marker number
- Origin_marker: Marker origin (e.g., nuclear, mtDNA)
- Type_marker: Marker type (e.g., SNP, microsatellite)
- Markers_stated_diagnostic: Whether markers are diagnostic (yes/no)
- Clines_between_0.2_0.8: Indicator if cline spans 0.2–0.8 frequency/ancestry range
- Cline_based_on: Data basis (e.g., allele frequency, ancestry)
- Cline_width_km: Width of cline (km)
- Excluded: Whether excluded from analysis (yes/no)
- Cline_to_use: Indicator for inclusion in final analyses
- Exclude_double_cline: Flag for duplicate clines
- Reason_for_exclusion: Explanation for exclusion
Sheet: All_transects
Description: Detailed summary of transect-level measurements.
Variables (selected):
- Transect_ID: Unique transect identifier.
- Source_ID: Identifier for the source study.
- number of single mitochondrial cline widths: Number of mitochondrial cline-width estimates.
- median mtDNA cline width [km]: Median mitochondrial cline width in kilometers.
- Lower support limits of mtDNA cline width [km]: Lower confidence bound of mitochondrial cline width.
- Upper support limits of mtDNA cline width [km]: Upper confidence bound of mitochondrial cline width.
- single locus nuclear cline widths: Indicator of availability of single-locus nuclear clines.
- number of single-locus nuclear cline widths: Number of single-locus nuclear cline estimates.
- median of single nuclear cline widths [km]: Median nuclear cline width in kilometers.
- multiple loci nuclear cline width: Indicator of availability of multilocus nuclear cline.
- number of loci used for nuclear genome average (Q ancestries) cline width: Number of loci used for genome-wide ancestry estimates.
- nuclear genome average (Q ancestries) cline width [km]: Genome-wide cline width estimate in kilometers.
- Lower support limits of nuclear genome average (Q ancestries) cline width [km]: Lower confidence bound for genome-wide cline width.
- Upper support limits of nuclear genome average (Q ancestries) cline width [km]: Upper confidence bound for genome-wide cline width.
- number of loci in a subset of SNPs used for second cline width: Number of loci used for alternative estimate.
- second cline width based on subset of loci or populations [km]: Alternative cline width estimate in kilometers.
Units: kilometers (km)
Sheet: All_cline_data
Description: Raw cline measurements extracted from literature.
Variables:
- Transect_ID: Transect identifier.
- HZ_ID: Hybrid zone identifier.
- Reference_ID: Source reference identifier.
- year: Publication year.
- details: Additional details describing the measurement.
- category (A = single nuclear cline, B = genome average, C = mtDNA): Type of cline estimate.
- type: Marker or data type (e.g., nuclear DNA, mtDNA, microsatellites).
- num_markers: Number of loci used.
- width (km): Cline width in kilometers.
- center (km): Cline center in kilometers.
- ρmin: Minimum allele frequency or ancestry value.
- ρmax: Maximum allele frequency or ancestry value.
- ρmax-ρmin: Range of allele frequency or ancestry values.
- method: Method used to estimate cline parameters.
- notes: Additional comments.
Sheet: Divergence_times
Description: Divergence time estimates for hybridizing taxa.
Variables:
- HZ_ID: Hybrid zone identifier.
- Taxon_1: First taxon.
- Taxon_2: Second taxon.
- div time estimate in lit (mya): Divergence time from literature (million years ago).
- div time estimate literatur to use (Mya): Selected literature-based estimate.
- divergence_time_reference: Source reference for divergence estimate.
- details: Additional notes or context.
- timetree estimated time (mya): TimeTree estimate.
- timetree estimated time range (mya): Range associated with TimeTree estimate.
- timetree div. times checked July 2022: Indicator of verification date.
- Median time (MYA): Median divergence time.
- Lower CI: Lower confidence interval.
- Upper CI: Upper confidence interval.
- Adjusted time: Standardized divergence time used for comparison.
- Number of cited studies: Number of studies contributing to estimate.
- Time to use: Final divergence time used in analyses.
Units: million years ago (mya)
Sheet: Dispersal_rates
Description: Dispersal estimates for taxa in hybrid zones.
Variables:
- HZ_ID: Hybrid zone identifier.
- Taxon_1: First taxon.
- Taxon_2: Second taxon.
- dispersal_category: Categorical grouping of dispersal values.
- dispersal_used: Selected dispersal value used in analyses.
- dispersal_(km/gen): Dispersal in kilometers per generation.
- dispersal_(km/sqrt(gen)): Dispersal scaled by the square root of generations.
- dispersal_(other): Dispersal reported in alternative units.
- other_dispersal_units: Units corresponding to alternative dispersal values.
- generation_time_(years/gen): Generation time in years per generation.
- dispersal_reference: Source reference for dispersal estimate.
- details: Additional notes or context.
- taxonomic_precision: Taxonomic level at which the estimate applies.
Units:
- km/gen = kilometers per generation
- km/sqrt(gen) = dispersal scaled by square root of generations
- years/gen = years per generation
Sheet: Input_for_R
Description: Curated dataset used as input for R analyses.
Variables: A subset and integration of cline widths, divergence estimates, and dispersal values used for modeling and statistical analysis.
Directory: TInflA/
Description: This directory contains all input data, phylogenetic trees, and results from Taxonomic Inflation Analyses (TInflA). These analyses quantify phylogenetic divergence and evaluate species delimitation patterns across taxonomic groups.
The directory is structured into three main subfolders:
- results/
- species_lists/
- timetrees/
Subdirectory: results/
Description: Contains output files from TInflA analyses. Each subfolder corresponds to one taxonomic group and analysis configuration.
Folder naming convention
Each folder follows the format:
TInflA_[TaxonomicGroup]adjusted_timetree_threshold[value]_[date]/
Examples:
- TInflA_Aves_adjusted_timetree_threshold_4_2022-06-01
- TInflA_Lissamphibia_adjusted_timetree_threshold_5.2_2023-02-13
Where:
- TaxonomicGroup = Aves, Mammalia, Squamata, etc.
- threshold = patristic distance cutoff used to define species boundaries
- date = date of analysis run
Contents of each results subfolder
Each folder contains paired files:
- ####_out.txt
- ####_tree.txt
Where:
- #### is the cut-off year used in the TInflA analysis
Each file, therefore, represents results based on species described up to that year.
File: ####_out.txt
Description: Primary output file listing species pairs and their phylogenetic (patristic) distances, along with summary statistics for a given cut-off year.
Variables:
- Species: Name of focal species
- Sister taxa: Closest related species in the phylogeny
- patristic distance: Phylogenetic distance between the species pair
Units:
- Patristic distance: branch length units derived from the time trees
File: ####_tree.txt
Description: Phylogenetic tree corresponding to the analysis for a given cut-off year.
Structure:
- First line: cut-off year (matching the filename)
- Remaining content: ASCII-formatted phylogenetic tree
Contents:
- Species names at tips
- Branching structure representing evolutionary relationships
Purpose:
- Used to calculate patristic distances reported in the corresponding ####_out.txt file
Subdirectory: species_lists/
Description: Contains species lists used as input for TInflA analyses.
File naming convention
TInflA_[TaxonomicGroup]specieslist_adjusted_timetree[date].tsv
Examples:
- TInflA_Aves_specieslist_adjusted_timetree_2022-06-01.tsv
- TInflA_Mammalia_specieslist_adjusted_timetree_2022-06-01.tsv
File contents
Each file contains:
- Species names included in the analysis
- Year of description
Subdirectory: timetrees/
Description: Contains phylogenetic trees used in TInflA analyses.
File naming convention
TInflA_[N][TaxonomicGroup][timetree_type]_[date].tre
Examples:
- TInflA_2339_Mollusca_adjusted_timetree_2022-06-01.tre
- TInflA_4417_Aves_adjusted_timetree_2022-06-01.tre
Where:
- N = number of taxa included in the tree
File contents
- Phylogenetic trees in Newick format
- Branch lengths represent evolutionary divergence time
Units:
- Branch lengths correspond to patristic distances used in analyses
Directory: R/
Description: This folder contains all scripts and input files used to generate figures (Figs. 1–3, S2–S10) and perform statistical analyses associated with the dataset.
File: R_Script_Figures_and_Statistics.R
Description: Main R script used to:
- Generate Figures 1–3 and Supplementary Figures S2–S10
- Perform all statistical analyses presented in the study
Inputs:
- HZ_data_input.txt
- TaxInf_input.txt
- specieslist.txt
- distances_time.txt
Outputs:
- Figures (plots)
- Statistical model results
File: HZ_data_input.txt
Description: Pre-filtered hybrid zone dataset used as the primary input for analyses. This file was exported from Hybrid_zone_data.xlsx and contains only the subset of data used in statistical models.
Variables:
- HZ_ID: Hybrid zone identifier
- Transect_ID: Transect identifier
- Hybridizing_taxa: Pair of taxa forming the hybrid zone
- Group: Taxonomic group
- Transect_Lit_ID: Literature-based transect identifier
- Number_marker: Number of genetic markers
- Number_marker_cat: Categorical bin of marker number
- Origin_marker: Marker origin (e.g., nuDNA, mtDNA)
- Type_marker: Marker type (e.g., SNPs, microsatellites)
- Markers_stated_diagnostic: Whether markers are diagnostic
- Clines_between_0.2_0.8: Indicator of cline shape coverage
- Cline_based_on: Basis for cline estimation
- Cline_width_km: Cline width (km)
- Lit_div_estimates: Divergence time from literature (mya)
- Timetree_div_estimates: Divergence time from TimeTree (mya)
- Dispersal_estimates_km/generation: Dispersal estimate (km/gen)
- Excluded: Exclusion flag (0 = included, 1 = excluded)
- Cline_to_use: Inclusion flag for analyses
- Exclude_double_cline: Flag for duplicate clines
Units:
- Distance: kilometers (km)
- Time: million years ago (mya)
- Dispersal: km per generation
File: distances_time.txt
Description: Contains results from all TInflA runs. Each row represents a species pair and their patristic distance.
Variables:
- Species_1: First species in the pair
- Species_2: Second species in the pair
- Patristic_dist: Phylogenetic (patristic) distance between species
- Group: Taxonomic group
Units:
- Patristic distance: branch length units from phylogenetic tree
File: specieslist.txt
Description: List of all species included in TInflA analyses, along with their taxonomic group and year of formal description.
Variables:
- Species: Scientific name (Genus_species format)
- Year: Year species was formally described
- Group: Taxonomic group
Units:
- Year: calendar year
File: TaxInf_input.txt
Description: Summary of patristic distances across time intervals, used to evaluate temporal patterns in species divergence.
Variables:
- time_interval: Year or cutoff used in analysis
- n_species_pairs: Number of species pairs included
- mean_patristic_distance: Mean phylogenetic distance
- minimum_patristic_distance: Minimum observed distance
- maximum_patristic_distance: Maximum observed distance
- pairs_under_threshold: Number of pairs below threshold
- perc_under_threshold: Proportion below threshold
- Group: Taxonomic group
- threshold: Distance threshold used
Units:
- Patristic distance: phylogenetic branch length units
- perc_under_threshold: proportion (0–1)
Missing Data and Use of “NA”
Missing values in the dataset are indicated as “NA”. These represent cases where data were not available from the original source, not reported in the literature, or not applicable for a given entry.
In some instances, values are missing because:
- The original study did not report the parameter (e.g., dispersal or divergence time)
- The parameter could not be reliably estimated
- The variable is not applicable to the specific marker type or analysis
Empty cells were replaced with “NA” where possible to minimize ambiguity.
Code/software
The software "TaxoInflationAnalyzer" or "TInflA" was used to generate files in the "TInflA" folder, which is described comprehensively on GitHub: https://github.com/iTaxoTools/TaxoInflationAnalyzer
All other files can be opened by a standard text file program, and the .xlsx file with Microsoft Excel.
All packages that were used are stated in the R script.
Access information
Data was derived from the following sources:
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