Are interphylum spiralian relationships resolvable?
Data files
Apr 16, 2026 version files 5.41 GB
-
Alignments.zip
9.53 MB
-
Astral.zip
502.92 KB
-
cladeRemoval.zip
901.31 MB
-
divTest.zip
1.45 GB
-
EDCfiles.zip
147.74 KB
-
guideTrees.zip
187.72 KB
-
IQTreeRuns.zip
702.02 MB
-
pbFiles.zip
12.96 MB
-
README.md
6.70 KB
-
Results.zip
206.49 MB
-
starTreeSims.zip
1.74 GB
-
taxonJack.zip
381.82 MB
Abstract
While the membership of the major animal clade of Spiralia has been relatively stable since the introduction of molecular sequence data, the relationships between the constituent phyla are less clear. Focusing on the five largest phyla (Annelida, Brachiopoda, Mollusca, Nemertea, and Platyhelminthes), we find previous analyses have supported all 15 possible unrooted trees that could relate them, suggesting a hard-to-resolve node. We have used two recent phylogenomic data sets to explore this remarkable example of taxonomic instability. Using a combination of taxon-jackknifing and bootstrapping on empirical and simulated data, we explored the support for all 105 rooted and 15 unrooted 5-phylum topologies under site-homogeneous and site-heterogeneous substitution models. Rooted analyses suggested that the preference for rooting Spiralia on Platyhelminthes is due to artefactual long-branch attraction between this clade and Gnathifera. Most unrooted analyses showed a marginal and non-significant preference for the same 5-phylum topology. In all unrooted trees, the branches relating to spiralian phyla were shorter, on average, than the short (and contested) Deuterostome branch. Our results suggest that spiralian phyla likely emerged in rapid succession, in a difficult to resolve adaptive radiation. Resolving these very short branches will require large data sets to overcome stochastic errors, as well as efforts to address systematic errors arising from both branch-length and site-compositional heterogeneity. The lack of clarity has implications for our understanding of the clade’s history, the interpretation of Cambrian fossils, and for clarifying the evolutionary history of traits, including biomineralization, segmentation, and larvae.
Directory list:
For all folders:
- MARL subfolders contain the data from and analyses based on Marlétaz et al. 2019
- SERR subfolders contain the data from and analyses based on Serra Silva et al. 2025.
- Any files with 'Pasc' in the name belong to this data set.
For all folders, except results.zip, the order of the 105 rooted topologies does not correspond to that depicted in Serra Silva and Telford 202x.
To covert between raw analyses and plots use this re-ordering: (99:105,8:14,15:21,1:7,22:28,29:35,36:42,43:49,50:56,57:63,64:70,71:77,78:84,85:91,92:98)
Alignments.zip folder: contains the concatenated alignments used for fixed-taxon topology-scoring analyses
- Concatenated:
- MARL:
- Marl_clanChecked_LophOgps_concat_70k.fasta: concatenated matrix for 105-rooted topology analyses
- Marl_clanChecked_LophOgps_concat_70k_5taxon.fasta: concatenated matrix for 15-unrooted topology analyses
- SERR:
- Pasc_loph_Outgps.fasta: concatenated matrix for 105-rooted topology analyses
- Pasc_noGnath_concat.fasta: concatenated matrix for 15-unrooted topology analyses
- MARL:
- Raw: locus alignments filtered to taxa of interest
Astral.zip: contains the input gene trees and inferred Astral trees for Marl and SERR data.
- Each MARL and SERR folders contain these subfolders:
- lociTrees: IQTREE runs for each locus
- STs: Astral-inferred trees with local posterior probabilities and quartet scores
cladeRemoval.zip: contains the files needed to generate and process analyses where Gnathifera or Platyhelminthes were removed from the matrices
- Alignments contains the concatenated fasta files organised by parent matrix (MARL or SERR)
- guideTrees contains the four sets of guide trees needed to repeat the analyses
*. RELL contains the script used to geenrate the RELL distributions, the R workspaces with the distributions described in the MS and the *.sitelh files from all topology-scoring analyses. For ALL analyses, the complete IQTREE output is provided for T1.
divTest.zip: contains the files needed to run amino acid diversity tests and the raw outputs of the analyses
EDCfiles.zip: contains the IQTREE-compatible EDM category file generated with the EDCluster package
guideTrees.zip: contains the guide trees used for the IQTREE topology-scoring analyses and for the data simulations
- Each MARL and SERR folders contain these subfolders:
- resolved: guide trees for topology-scoring
- 15unrooted: fixed topologies for unrooted analyses
- 105rooted: fixed topologies for rooted analyses
- rootedStarTrees: input for data simulations
- resolved: guide trees for topology-scoring
IQTreeRuns.zip: contains the output files from IQTREE's topology-scoring runs, organised by original data set, rooted or unrooted, and substitution model used (LG or EDM)
intraPhylumSplits.zip: contains the unconstrained analysis on the SERR matrix used to define intraphylum branching. MARL intraphylum branching based on original publication.
pbFiles.zip: contains the PhyloBayes runs used to generate the site-profiles used as input in EDCfiles.zip. *.run files not included due to large size but can be requested from the authors.
Results.zip: contains the scripts needed to process the IQTree outputs and the edited files with log-likelihoods, branch-lengths and pseudo-bootstraps
- branchLengths: contains the scripts needed to calculate branch lengths and plot them. For both MARL and SERR subfolders:
- EDM: contains the tree file (all.tre) with all scored and optimised trees under EDM+G4 and respective branch-lengths (*.txt)
- LG: contains the tree file (all.tre) with all scored and optimised trees under LG+G4 and respective branch-lengths (*.txt)
- pynt_*Data_ogps.py: data set-specific Python3 script to extract branch lengths
- plots_BLconcat_ogps_LGvEDM_*.R: data set-specific R script to plot the branch-lengths of interest
- RELL: contains the scripts to generate RELL distributions and the R workspaces with the distributions generated for the MS
- 15unrooted:
- 5taxon_wksp.RData: RELL distributions used for MS
- siteLnL_bootstrap_5taxon.R: R script to generate and process RELL distributions
- 105rooted:
- all_wksp.RData: RELL distributions used for MS
- siteLnL_bootstrap.R: R script to generate and process RELL distributions
- 15unrooted:
- starSims: log-likelihood scores of topology-scoring on simulated data and scripts to process and plot results. The 15unrooted and 105rooted folders include:
- starSim_lnl_Marl_EDM.txt
- starSim_lnl_Marl_EDMunderLG.txt
- starSim_lnl_Pasc_EDM.txt
- starSim_lnl_Pasc_EDMunderLG.txt
- starSimsVisualisation*.R : script to process the four text files listed above.
- taxJacks: log-likelihood scores of topology-scoring on taxon-jackknives and scripts to process and plot results. The 15unrooted and 105rooted folders include:
- EDM:
- Marl_taxJack_lnL_nohead.txt
- Pasc_taxJack_lnL_nohead.txt
- LG:
- Marl_taxJack_lnL_nohead.txt
- Pasc_taxJack_lnL_nohead.txt
- taxJackVisualisation*.R
- EDM:
starTreeSims.zip: contains the input data for generating the simulated alignments, the simulated data and one example IQTREE run for each rooting+data+model combination
- topoLnL
- EDM: contains one IQTREE topology-scoring run for data simulated and scored under EDM
- EDM_underLG: contains one IQTREE topology-scoring run for data simulated under EDM and scored under LG
- simulatedData:
- alns_EDM: EDM-simulated alignments
- EDCfiles: contains the IQTREE-compatible EDM category files for every tenth simulated alignments
- inputFiles: input for AliSim simulations
- IQTreeFiles_EDM: AliSim run
- pbFiles: PhyloBayes runs used to obtain the site-profiles for every tenth simulated alignment. *.run files omitted due to large sizes.
taxonJack.zip: contains the taxon jackknife replicates (alignments and pruned trees) and one example IQTREE run for each rooting+data+model combination
- input
- ancillary: contains all files needed to generate the taxon-jackknives. See Serra Silva et al. 2025 data for details.
- Align: taxon-jackknifed alignments
- Trees: pruned trees fro each jackknife replicate
- topoLnL
- EDM: contains one IQTREE topology-scoring run under the EDM model
- LG: contains one IQTREE topology-scoring run under the LG model
