Are interphylum spiralian relationships resolvable?
Figures
There are 15 unrooted trees linking the 5 major spiralian phyla and each has been supported in at least one publication (Supplementary file 1).
Tree numbering throughout this study refers to the trees in this figure. For all 105 rooted trees, the rooting schemes follow that depicted in unrooted T1.
Branches linking spiralian phyla are shorter than both the short, disputed deuterostome branch (Kapli et al., 2021; Serra Silva et al., 2025) and the short branch leading to Lophotrochozoa (Telford et al., 2015).
(A and B) are data from SERR. (C and D) are data from MARL. Compared to the site-homogeneous model (A, C), internal branches (orange) become shorter under the site-heterogeneous model while the long branches leading to Platyhelminthes (blue) and Mollusca (dark green) become even longer (B, D). Circles on the y-axis indicate the median branch lengths of Deuterostomia (D), Ecdysozoa (E), Lophotrochozoa (L), and Protostomia (P). Triangles correspond to the median spiralian internal branches (Int), values given in Appendix 2—table 1. Scales are identical on either side of the y-axis breaks.
Discordance-aware summary tree analyses show extensive gene-tree conflict within Spiralia.
For SERR (A), the Astral analysis recovers all spiralian phyla as monophyletic but no internal Spiralia branch with quartet scores (QS) over 0.50. The exception is the branch separating Platyhelminthes + outgroups from the remaining Spiralia phyla; however, Platyhelminthes are recovered within paraphyletic Gnathifera. For MARL (B), the Astral analysis yields very similar results, except that Annelida is not monophyletic. Branch-lengths are in coalescent units and monophyletic phyla have been collapsed, with the QS (Q1/Q2/Q3) for the branch subtending each phylum listed underneath the phylum names. Detailed QS can be found in the Astral.zip folder in the Dryad data repository (https://doi.org/10.5061/dryad.280gb5n3j).
Relative median resampling estimated log-likelihood (RELL) pseudo-bootstraps (A–D) and randomised taxon-jackknives (E–H) log-likelihood ranks for all rooted trees, see Appendix 2—table 3 and Appendix 2—table 5 for all ranks.
Highest- and lowest-ranked topologies labelled for all analyses, along with every seventh topology from highest- to lowest-ranked. Topologies ranked by their difference in log-likelihood from the mean log-likelihood of each RELL or jackknife distribution. The RELL analyses of both datasets under the site-homogeneous LG + F + G4 model (A, B) occupy the widest ranges of log-likelihoods, which is consistent with the statistically significant differences found by the Kruskal–Wallis (KW) test. The RELL analyses under the site-heterogeneous EDM + F + G4 model (C, D) occupy a much narrower range of log-likelihoods. In practical terms, the wider the log-likelihood range, the more dissimilar scores are and the more we would expect to recover significantly different scores between topologies. The jackknife analyses show a similar trend, wider range of median log-likelihood scores under LG + F + G4 (E, F) than under EDM + F + G4 (G, H); however, the log-likelihood ranges of the jackknives under LG + F + G4 are very similar to that occupied by RELL under EDM + F + G4, consistent with the lack of statistically significant differences between topologies in the jackknife analyses.
With simulated data inadequate models incorrectly support a platyhelminth root.
Using data simulated on an unresolved Spiralia tree, we ranked rooted spiralian topologies by difference to mean log-likelihood score. Highest- and lowest-ranked topologies are labelled for all analyses, along with every seventh-topology from highest- to lowest-ranked. For datasets simulated under EDM (complex) but scored under LG (simple) the 15 topologies with Spiralia rooted on Platyhelminthes are highest ranked (A, B). The plots for the ‘correctly’ specified models (C, D) show the expected approximately equal log-likelihood scores for all 105 rooted topologies. See Appendix 2—table 7 for complete topology rankings.
Most fixed-taxon resampling estimated log-likelihood (RELL) bootstrap and randomised taxon-jackknife replicates, under both site-homogeneous (LG) and site-heterogeneous (EDM) models, recover unrooted T4 as the best-scoring topology.
Heights of bars correspond to the number of replicates where each topology had the highest log-likelihood score.
Relative median resampling estimated log-likelihood (RELL) pseudo-bootstraps (A–D) and randomised taxon-jackknives (E–H) log-likelihood ranks for all unrooted trees, see Appendix 2—table 8 and Appendix 2—table 10 for all ranks.
Topologies ranked by their difference in log-likelihood to the mean log-likelihood of each RELL or jackknife distribution. The RELL analyses of both datasets under the site-homogeneous LG + F + G4 model (A, B) occupy the widest ranges of log-likelihoods, which is consistent with the statistically significant differences found by the Kruskal-Wallis test. The RELL analyses under the site-heterogeneous EDM + F + G4 model (C, D) occupy a much narrower range of log-likelihoods. The jackknife analyses show a similar trend, wider range of median log-likelihood scores under LG + F + G4 (E, F) than under EDM + F + G4 (G, H); however, the log-likelihood ranges of the jackknives under LG + F + G4 are very similar to that occupied by RELL under EDM + F + G4, consistent with the lack of statistically significant differences between topologies in the jackknife analyses.
With simulated data, misspecified models recover unrooted T4 within the set of highest-scoring topologies.
Using data simulated on an unresolved Spiralia tree we ranked unrooted spiralian topologies by difference to mean log-likelihood score. For datasets simulated under EDM (complex) but scored under LG (simple), three topologies (uT4, uT5, and uT14) are within the highest-ranked topologies for both empirical datasets (A, B). The plots for the ‘correctly’ specified models (C, D) show the expected approximately equal log-likelihood scores for all 15 unrooted topologies.
Tables
| Reagent type (species) or resource | Designation | Source or reference | Identifiers | Additional information |
|---|---|---|---|---|
| Other | Alignments, phylogenetic analyses, simulations, and ancillary scripts | DRYAD; https://doi.org/10.5061/dryad.280gb5n3j | Please see Materials and methods for data processing details | |
| Software and algorithms | IQ-TREE2 (v.2.3.5) | Minh et al., 2020; http://www.iqtree.org/ | ||
| Software and algorithms | EDCluster | Schrempf et al., 2020; Schrempf, 2020; https://github.com/dschrempf/EDCluster | ||
| Software and algorithms | ASTRAL-III (v.5.1.3) | Zhang et al., 2017a; Zhang et al., 2017b; https://github.com/smirarab/ASTRAL | ||
| Software and algorithms | MAFFT (v.7.455) | Katoh and Standley, 2013; https://mafft.cbrc.jp/alignment/software/ | ||
| Software and algorithms | trimAl (v.1.4.rev15) | Capella-Gutiérrez et al., 2009; https://vicfero.github.io/trimal/ | ||
| Software and algorithms | Amino acid diversity tests | Giacomelli et al., 2022; Giacomelli et al., 2025; Giacomelli, 2026; https://github.com/mgiacom/tardigrades_catpmsf | ||
| Software and algorithms | Additional data processing and analysis scripts | DRYAD; https://doi.org/10.5061/dryad.280gb5n3j |
Median branch lengths for the 105 rooted topologies.
Phylum-level branch lengths shown in Figure 2.
| Branch | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| Deuterostomia | 0.03139 | 0.02812 | 0.02305 | 0.01782 |
| Ecdysozoa | 0.04762 | 0.06257 | 0.03835 | 0.03941 |
| Lophotrochozoa | 0.02440 | 0.02607 | 0.02382 | 0.02613 |
| Protostomia | 0.06420 | 0.09325 | 0.04852 | 0.06641 |
| Internal | 0.01616 | 0.01393 | 0.01316 | 0.009462 |
Fixed-taxon RELL pseudo-bootstraps rankings for 105 rooted topologies.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 1 | 40 | 34 | 26 | 12 |
| 2 | 5 | 3 | 2 | 8 |
| 3 | 38 | 35 | 15 | 7 |
| 4 | 73 | 77 | 46 | 27 |
| 5 | 71 | 57 | 38 | 21 |
| 6 | 92 | 81 | 77 | 77 |
| 7 | 101 | 91 | 61 | 80 |
| 8 | 56 | 66 | 80 | 24 |
| 9 | 35 | 31 | 48 | 6 |
| 10 | 27 | 22 | 40 | 11 |
| 11 | 41 | 42 | 56 | 62 |
| 12 | 19 | 21 | 51 | 55 |
| 13 | 30 | 24 | 66 | 57 |
| 14 | 2 | 2 | 13 | 40 |
| 15 | 72 | 70 | 68 | 23 |
| 16 | 50 | 36 | 39 | 5 |
| 17 | 33 | 23 | 34 | 9 |
| 18 | 4 | 1 | 6 | 10 |
| 19 | 43 | 25 | 42 | 19 |
| 20 | 77 | 54 | 70 | 71 |
| 21 | 64 | 45 | 82 | 67 |
| 22 | 39 | 38 | 37 | 4 |
| 23 | 22 | 18 | 22 | 1 |
| 24 | 17 | 12 | 20 | 2 |
| 25 | 21 | 16 | 32 | 18 |
| 26 | 18 | 13 | 12 | 15 |
| 27 | 45 | 26 | 35 | 49 |
| 28 | 6 | 4 | 5 | 29 |
| 29 | 57 | 62 | 53 | 33 |
| 30 | 46 | 47 | 52 | 37 |
| 31 | 34 | 28 | 25 | 14 |
| 32 | 23 | 19 | 14 | 3 |
| 33 | 20 | 14 | 8 | 13 |
| 34 | 55 | 33 | 29 | 51 |
| 35 | 10 | 5 | 3 | 30 |
| 36 | 97 | 96 | 69 | 59 |
| 37 | 90 | 86 | 71 | 61 |
| 38 | 75 | 60 | 33 | 31 |
| 39 | 78 | 68 | 31 | 52 |
| 40 | 48 | 40 | 10 | 20 |
| 41 | 49 | 39 | 21 | 26 |
| 42 | 7 | 6 | 1 | 22 |
| 43 | 100 | 92 | 95 | 72 |
| 44 | 89 | 78 | 101 | 79 |
| 45 | 70 | 53 | 49 | 43 |
| 46 | 13 | 8 | 9 | 28 |
| 47 | 74 | 51 | 59 | 53 |
| 48 | 93 | 69 | 65 | 68 |
| 49 | 67 | 46 | 41 | 32 |
| 50 | 82 | 102 | 99 | 99 |
| 51 | 81 | 100 | 85 | 103 |
| 52 | 47 | 88 | 67 | 87 |
| 53 | 29 | 49 | 45 | 44 |
| 54 | 16 | 32 | 58 | 83 |
| 55 | 26 | 43 | 72 | 88 |
| 56 | 1 | 7 | 19 | 66 |
| 57 | 102 | 104 | 73 | 86 |
| 58 | 103 | 105 | 60 | 94 |
| 59 | 66 | 95 | 44 | 69 |
| 60 | 52 | 75 | 50 | 70 |
| 61 | 36 | 55 | 16 | 34 |
| 62 | 37 | 52 | 28 | 42 |
| 63 | 3 | 9 | 4 | 35 |
| 64 | 98 | 103 | 96 | 101 |
| 65 | 96 | 101 | 86 | 104 |
| 66 | 60 | 90 | 64 | 90 |
| 67 | 8 | 20 | 11 | 65 |
| 68 | 54 | 89 | 63 | 84 |
| 69 | 51 | 63 | 55 | 46 |
| 70 | 59 | 76 | 90 | 91 |
| 71 | 65 | 64 | 74 | 41 |
| 72 | 11 | 11 | 17 | 25 |
| 73 | 63 | 61 | 79 | 45 |
| 74 | 58 | 44 | 54 | 17 |
| 75 | 84 | 59 | 75 | 38 |
| 76 | 99 | 82 | 105 | 98 |
| 77 | 105 | 93 | 102 | 102 |
| 78 | 88 | 83 | 94 | 74 |
| 79 | 14 | 17 | 27 | 58 |
| 80 | 85 | 79 | 100 | 82 |
| 81 | 91 | 80 | 103 | 93 |
| 82 | 94 | 71 | 97 | 76 |
| 83 | 86 | 65 | 78 | 56 |
| 84 | 104 | 87 | 98 | 96 |
| 85 | 87 | 98 | 89 | 95 |
| 86 | 15 | 27 | 24 | 78 |
| 87 | 83 | 97 | 92 | 100 |
| 88 | 95 | 99 | 88 | 105 |
| 89 | 80 | 94 | 87 | 92 |
| 90 | 69 | 72 | 81 | 54 |
| 91 | 79 | 84 | 104 | 97 |
| 92 | 25 | 29 | 30 | 16 |
| 93 | 31 | 41 | 57 | 47 |
| 94 | 32 | 56 | 36 | 39 |
| 95 | 42 | 74 | 43 | 48 |
| 96 | 24 | 30 | 18 | 36 |
| 97 | 62 | 67 | 47 | 85 |
| 98 | 12 | 15 | 7 | 63 |
| 99 | 53 | 50 | 62 | 50 |
| 100 | 76 | 73 | 84 | 89 |
| 101 | 61 | 58 | 83 | 64 |
| 102 | 68 | 85 | 93 | 75 |
| 103 | 28 | 37 | 76 | 73 |
| 104 | 44 | 48 | 91 | 81 |
| 105 | 9 | 10 | 23 | 60 |
Fixed-taxon RELL pseudo-bootstraps best-ranked topology frequencies for 105 rooted topologies.
Omitted topologies were never recovered as the best-ranked tree.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 2 | 8.4 | 11.96 | 27.62 | 0.43 |
| 3 | 0 | 0 | 0 | 0.59 |
| 14 | 8.09 | 15.74 | 0 | 0 |
| 18 | 11.23 | 51.78 | 1.72 | 0.34 |
| 22 | 0 | 0 | 0 | 0.04 |
| 23 | 0 | 0 | 0 | 57.2 |
| 24 | 0 | 0.28 | 0 | 36.52 |
| 26 | 0 | 0.1 | 0.01 | 0.05 |
| 28 | 0.41 | 3.8 | 4.72 | 0 |
| 29 | 0 | 0 | 0 | 0.01 |
| 31 | 0 | 0 | 0 | 0.67 |
| 32 | 0 | 0 | 0 | 3.53 |
| 33 | 0 | 0.04 | 0.36 | 0.5 |
| 35 | 0.26 | 6.15 | 13.15 | 0 |
| 38 | 0 | 0 | 0 | 0.01 |
| 39 | 0 | 0 | 0 | 0 |
| 40 | 0 | 0 | 0 | 0.01 |
| 42 | 0.34 | 5.26 | 50.33 | 0.05 |
| 46 | 0.01 | 1.49 | 0 | 0.02 |
| 56 | 63.53 | 2.23 | 0 | 0 |
| 61 | 0 | 0 | 0 | 0.02 |
| 63 | 7.65 | 0.54 | 2.03 | 0 |
| 67 | 0.03 | 0 | 0 | 0 |
| 72 | 0 | 0.07 | 0 | 0 |
| 92 | 0 | 0 | 0 | 0.01 |
| 98 | 0 | 0 | 0.06 | 0 |
| 105 | 0.05 | 0.5 | 0 | 0 |
Randomised taxon-jackknife rankings for 105 rooted topologies.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 1 | 39 | 18 | 36 | 19 |
| 2 | 7 | 15 | 3 | 24 |
| 3 | 22 | 14 | 6 | 10 |
| 4 | 63 | 64 | 30 | 13 |
| 5 | 54 | 13 | 28 | 17 |
| 6 | 80 | 54 | 75 | 83 |
| 7 | 100 | 88 | 65 | 72 |
| 8 | 58 | 45 | 85 | 11 |
| 9 | 33 | 12 | 52 | 6 |
| 10 | 20 | 2 | 35 | 4 |
| 11 | 48 | 41 | 50 | 53 |
| 12 | 15 | 7 | 45 | 50 |
| 13 | 34 | 16 | 77 | 69 |
| 14 | 4 | 9 | 19 | 70 |
| 15 | 77 | 58 | 71 | 12 |
| 16 | 52 | 11 | 48 | 7 |
| 17 | 24 | 4 | 26 | 5 |
| 18 | 2 | 3 | 1 | 14 |
| 19 | 35 | 5 | 43 | 20 |
| 20 | 91 | 61 | 79 | 76 |
| 21 | 67 | 27 | 91 | 77 |
| 22 | 55 | 33 | 38 | 3 |
| 23 | 38 | 6 | 22 | 2 |
| 24 | 19 | 1 | 11 | 1 |
| 25 | 27 | 8 | 31 | 23 |
| 26 | 23 | 10 | 9 | 16 |
| 27 | 56 | 37 | 47 | 60 |
| 28 | 9 | 24 | 12 | 54 |
| 29 | 72 | 59 | 63 | 26 |
| 30 | 61 | 40 | 64 | 40 |
| 31 | 43 | 22 | 20 | 9 |
| 32 | 31 | 31 | 15 | 8 |
| 33 | 36 | 65 | 10 | 18 |
| 34 | 75 | 87 | 44 | 67 |
| 35 | 13 | 80 | 13 | 58 |
| 36 | 93 | 81 | 69 | 46 |
| 37 | 79 | 63 | 78 | 63 |
| 38 | 64 | 55 | 25 | 25 |
| 39 | 92 | 92 | 27 | 42 |
| 40 | 30 | 56 | 7 | 22 |
| 41 | 42 | 46 | 32 | 28 |
| 42 | 8 | 42 | 4 | 37 |
| 43 | 96 | 76 | 96 | 71 |
| 44 | 90 | 70 | 102 | 90 |
| 45 | 60 | 38 | 58 | 44 |
| 46 | 11 | 26 | 16 | 52 |
| 47 | 68 | 47 | 67 | 57 |
| 48 | 101 | 93 | 72 | 78 |
| 49 | 69 | 53 | 57 | 30 |
| 50 | 82 | 100 | 99 | 85 |
| 51 | 83 | 104 | 86 | 94 |
| 52 | 29 | 69 | 54 | 56 |
| 53 | 21 | 21 | 39 | 32 |
| 54 | 12 | 17 | 49 | 66 |
| 55 | 28 | 30 | 76 | 82 |
| 56 | 1 | 20 | 17 | 84 |
| 57 | 95 | 101 | 60 | 61 |
| 58 | 99 | 105 | 55 | 75 |
| 59 | 46 | 77 | 23 | 38 |
| 60 | 37 | 50 | 42 | 55 |
| 61 | 16 | 39 | 5 | 33 |
| 62 | 26 | 34 | 24 | 43 |
| 63 | 3 | 32 | 2 | 50 |
| 64 | 97 | 98 | 92 | 87 |
| 65 | 102 | 102 | 88 | 95 |
| 66 | 44 | 71 | 46 | 68 |
| 67 | 6 | 60 | 8 | 65 |
| 68 | 45 | 68 | 51 | 64 |
| 69 | 50 | 35 | 56 | 39 |
| 70 | 49 | 43 | 90 | 89 |
| 71 | 70 | 75 | 70 | 34 |
| 72 | 10 | 57 | 21 | 41 |
| 73 | 66 | 79 | 61 | 45 |
| 74 | 53 | 25 | 53 | 21 |
| 75 | 71 | 23 | 62 | 36 |
| 76 | 98 | 62 | 104 | 105 |
| 77 | 105 | 89 | 100 | 102 |
| 78 | 87 | 84 | 98 | 74 |
| 79 | 14 | 78 | 41 | 91 |
| 80 | 81 | 86 | 93 | 93 |
| 81 | 84 | 74 | 103 | 104 |
| 82 | 88 | 66 | 95 | 79 |
| 83 | 86 | 72 | 80 | 49 |
| 84 | 104 | 95 | 97 | 92 |
| 85 | 89 | 97 | 87 | 96 |
| 86 | 17 | 94 | 34 | 100 |
| 87 | 85 | 99 | 82 | 101 |
| 88 | 103 | 103 | 84 | 103 |
| 89 | 73 | 83 | 73 | 81 |
| 90 | 76 | 44 | 81 | 48 |
| 91 | 78 | 52 | 105 | 99 |
| 92 | 40 | 19 | 33 | 15 |
| 93 | 47 | 28 | 66 | 47 |
| 94 | 41 | 49 | 29 | 29 |
| 95 | 62 | 85 | 40 | 31 |
| 96 | 32 | 73 | 14 | 27 |
| 97 | 74 | 96 | 59 | 88 |
| 98 | 18 | 90 | 18 | 86 |
| 99 | 57 | 67 | 68 | 35 |
| 100 | 94 | 91 | 89 | 73 |
| 101 | 59 | 51 | 83 | 59 |
| 102 | 65 | 82 | 94 | 62 |
| 103 | 25 | 29 | 74 | 80 |
| 104 | 51 | 48 | 101 | 98 |
| 105 | 5 | 36 | 37 | 97 |
Randomised taxon-jackknife best-ranked topology frequencies for 105 rooted topologies.
Omitted topologies were never recovered as the best-ranked tree.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 2 | 15 | 0 | 0 | 0 |
| 8 | 0 | 0 | 0 | 5 |
| 9 | 0 | 0 | 0 | 0 |
| 10 | 0 | 10 | 0 | 5 |
| 12 | 0 | 5 | 0 | 0 |
| 14 | 5 | 5 | 0 | 0 |
| 17 | 0 | 5 | 0 | 0 |
| 18 | 30 | 5 | 25 | 0 |
| 23 | 0 | 5 | 0 | 30 |
| 24 | 0 | 40 | 0 | 55 |
| 26 | 0 | 5 | 5 | 0 |
| 31 | 0 | 5 | 0 | 0 |
| 32 | 0 | 5 | 0 | 5 |
| 33 | 0 | 0 | 15 | 0 |
| 40 | 0 | 0 | 10 | 0 |
| 42 | 0 | 0 | 5 | 0 |
| 54 | 10 | 0 | 0 | 0 |
| 56 | 30 | 0 | 0 | 0 |
| 61 | 0 | 0 | 10 | 0 |
| 63 | 10 | 0 | 20 | 0 |
| 67 | 0 | 0 | 10 | 0 |
| 92 | 0 | 5 | 0 | 0 |
| 103 | 0 | 5 | 0 | 0 |
Simulations on rooted Spiralia star-tree rankings for 105 rooted topologies.
For columns SERR + EDM and MARL +EDM, proportions are based on 10 alignments.
| Topology | SERR + LG EDM sim | SERR + EDM | MARL + LG EDM sim | MARL + EDM |
|---|---|---|---|---|
| 1 | 24 | 9 | 24 | 71 |
| 2 | 4 | 2 | 4 | 7 |
| 3 | 21 | 41 | 18 | 34 |
| 4 | 57 | 44 | 37 | 44 |
| 5 | 54 | 89 | 36 | 46 |
| 6 | 69 | 103 | 70 | 47 |
| 7 | 81 | 91 | 56 | 61 |
| 8 | 95 | 86 | 105 | 93 |
| 9 | 64 | 90 | 101 | 78 |
| 10 | 49 | 46 | 93 | 80 |
| 11 | 51 | 61 | 86 | 87 |
| 12 | 23 | 19 | 58 | 56 |
| 13 | 26 | 53 | 73 | 86 |
| 14 | 6 | 5 | 14 | 8 |
| 15 | 94 | 99 | 81 | 99 |
| 16 | 68 | 57 | 77 | 92 |
| 17 | 47 | 95 | 55 | 102 |
| 18 | 10 | 31 | 9 | 11 |
| 19 | 48 | 97 | 48 | 100 |
| 20 | 73 | 77 | 88 | 98 |
| 21 | 62 | 73 | 95 | 96 |
| 22 | 103 | 81 | 74 | 21 |
| 23 | 78 | 87 | 68 | 28 |
| 24 | 55 | 8 | 49 | 24 |
| 25 | 53 | 22 | 50 | 9 |
| 26 | 37 | 3 | 21 | 32 |
| 27 | 39 | 42 | 33 | 101 |
| 28 | 11 | 47 | 6 | 16 |
| 29 | 102 | 29 | 65 | 23 |
| 30 | 90 | 58 | 66 | 20 |
| 31 | 50 | 71 | 29 | 25 |
| 32 | 46 | 70 | 32 | 35 |
| 33 | 34 | 50 | 17 | 48 |
| 34 | 38 | 79 | 23 | 105 |
| 35 | 8 | 48 | 2 | 19 |
| 36 | 87 | 65 | 57 | 42 |
| 37 | 67 | 26 | 62 | 27 |
| 38 | 41 | 83 | 27 | 50 |
| 39 | 43 | 76 | 30 | 55 |
| 40 | 19 | 68 | 16 | 37 |
| 41 | 22 | 40 | 22 | 82 |
| 42 | 3 | 7 | 1 | 1 |
| 43 | 96 | 62 | 98 | 75 |
| 44 | 79 | 45 | 99 | 60 |
| 45 | 42 | 78 | 39 | 79 |
| 46 | 9 | 60 | 7 | 13 |
| 47 | 44 | 98 | 45 | 90 |
| 48 | 61 | 94 | 63 | 103 |
| 49 | 52 | 80 | 61 | 76 |
| 50 | 91 | 34 | 104 | 64 |
| 51 | 85 | 67 | 103 | 72 |
| 52 | 31 | 20 | 84 | 67 |
| 53 | 30 | 35 | 78 | 77 |
| 54 | 17 | 10 | 53 | 69 |
| 55 | 20 | 39 | 67 | 65 |
| 56 | 2 | 4 | 13 | 12 |
| 57 | 80 | 36 | 71 | 51 |
| 58 | 74 | 75 | 64 | 59 |
| 59 | 29 | 16 | 43 | 54 |
| 60 | 28 | 24 | 44 | 41 |
| 61 | 16 | 37 | 19 | 45 |
| 62 | 18 | 13 | 26 | 81 |
| 63 | 1 | 1 | 3 | 2 |
| 64 | 99 | 28 | 87 | 95 |
| 65 | 98 | 84 | 85 | 97 |
| 66 | 32 | 12 | 54 | 83 |
| 67 | 5 | 11 | 10 | 18 |
| 68 | 33 | 82 | 47 | 94 |
| 69 | 35 | 43 | 83 | 88 |
| 70 | 36 | 30 | 92 | 85 |
| 71 | 60 | 100 | 35 | 84 |
| 72 | 13 | 105 | 8 | 3 |
| 73 | 56 | 21 | 25 | 43 |
| 74 | 66 | 14 | 51 | 53 |
| 75 | 82 | 23 | 46 | 49 |
| 76 | 100 | 69 | 91 | 29 |
| 77 | 105 | 74 | 82 | 70 |
| 78 | 63 | 88 | 42 | 74 |
| 79 | 14 | 63 | 12 | 6 |
| 80 | 59 | 59 | 34 | 36 |
| 81 | 76 | 25 | 80 | 15 |
| 82 | 84 | 64 | 76 | 39 |
| 83 | 92 | 52 | 90 | 58 |
| 84 | 104 | 85 | 94 | 68 |
| 85 | 72 | 96 | 38 | 89 |
| 86 | 15 | 101 | 11 | 5 |
| 87 | 70 | 56 | 28 | 40 |
| 88 | 101 | 51 | 59 | 63 |
| 89 | 89 | 15 | 52 | 38 |
| 90 | 86 | 38 | 79 | 33 |
| 91 | 93 | 33 | 89 | 30 |
| 92 | 75 | 72 | 60 | 14 |
| 93 | 83 | 17 | 72 | 10 |
| 94 | 77 | 32 | 40 | 17 |
| 95 | 97 | 49 | 41 | 26 |
| 96 | 40 | 18 | 20 | 52 |
| 97 | 45 | 93 | 31 | 104 |
| 98 | 12 | 66 | 5 | 22 |
| 99 | 65 | 92 | 100 | 73 |
| 100 | 88 | 102 | 102 | 91 |
| 101 | 58 | 104 | 96 | 66 |
| 102 | 71 | 55 | 97 | 57 |
| 103 | 25 | 27 | 69 | 31 |
| 104 | 27 | 54 | 75 | 62 |
| 105 | 7 | 6 | 15 | 4 |
Simulations on rooted Spiralia star-tree best-ranked topology frequencies for 105 rooted topologies.
For columns SERR + EDM and MARL + EDM, proportions are based on 10 alignments. Omitted topologies were never recovered as the best-ranked tree.
| Topology | SERR + LG EDM sim | SERR + EDM | MARL + LG EDM sim | MARL + EDM |
|---|---|---|---|---|
| 10 | 0 | 0 | 0 | 20 |
| 18 | 0 | 20 | 0 | 0 |
| 23 | 0 | 20 | 0 | 0 |
| 28 | 0 | 0 | 15 | 0 |
| 33 | 0 | 0 | 1 | 0 |
| 35 | 0 | 0 | 84 | 0 |
| 49 | 2 | 0 | 0 | 0 |
| 56 | 98 | 0 | 0 | 0 |
| 60 | 0 | 20 | 0 | 0 |
| 72 | 0 | 0 | 0 | 20 |
| 85 | 0 | 0 | 0 | 20 |
| 86 | 0 | 0 | 0 | 20 |
| 89 | 0 | 20 | 0 | 0 |
| 91 | 0 | 20 | 0 | 0 |
| 98 | 0 | 0 | 0 | 20 |
Fixed-taxon RELL pseudo-bootstraps best-ranked topology frequencies for 15 unrooted topologies.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 1 | 0 | 0 | 0 | 0 |
| 2 | 19 | 2 | 0 | 3 |
| 3 | 0 | 0 | 0 | 0 |
| 4 | 81 | 98 | 60 | 96 |
| 5 | 0 | 0 | 35 | 1 |
| 6 | 0 | 0 | 5 | 0 |
| 7 | 0 | 0 | 0 | 0 |
| 8 | 0 | 0 | 0 | 0 |
| 9 | 0 | 0 | 0 | 0 |
| 10 | 0 | 0 | 0 | 0 |
| 11 | 0 | 0 | 0 | 0 |
| 12 | 0 | 0 | 0 | 0 |
| 13 | 0 | 0 | 0 | 0 |
| 14 | 0 | 0 | 0 | 0 |
| 15 | 0 | 0 | 0 | 0 |
Compositional diversity model adequacy tests on the fixed-taxon data matrices.
Data simulated under the EDM + G4 model are closest in amino acid compositional diversity to the empirical data. Bold values correspond to the simulated data that most closely resemble the empirical data.
| Model | Amino acid diversity | Z-score | |
|---|---|---|---|
| Empirical data | Simulated data | ||
| MARL | |||
| LG + G4 | 5.1592 | 5.4859 ± 0.005859 | 55.7581 |
| EDM + G4 | 5.1592 | 5.3082 ± 0.005771 | 25.8182 |
| SERR | |||
| LG + G4 | 6.2950 | 6.5769 ± 0.007286 | 38.6836 |
| EDM + G4 | 6.2950 | 6.1735 ± 0.006021 | –20.1752 |
Randomised taxon-jackknife best-ranked topology frequencies for 15 unrooted topologies.
| Topology | SERR + LG | SERR + EDM | MARL + LG | MARL + EDM |
|---|---|---|---|---|
| 1 | 2 | 0 | 4 | 0 |
| 2 | 23 | 11 | 0 | 2 |
| 3 | 0 | 0 | 0 | 2 |
| 4 | 38 | 61 | 43 | 92 |
| 5 | 0 | 6 | 26 | 1 |
| 6 | 0 | 0 | 12 | 0 |
| 7 | 0 | 0 | 0 | 0 |
| 8 | 30 | 0 | 0 | 1 |
| 9 | 5 | 1 | 15 | 1 |
| 10 | 0 | 0 | 0 | 0 |
| 11 | 0 | 0 | 0 | 0 |
| 12 | 0 | 0 | 0 | 0 |
| 13 | 0 | 0 | 0 | 0 |
| 14 | 2 | 20 | 0 | 1 |
| 15 | 0 | 1 | 0 | 0 |
Simulations on 5-taxon Spiralia star-tree best-ranked topology frequencies for 15 unrooted topologies.
For columns SERR + EDM and MARL + EDM, proportions are based on 10 alignments.
| Topology | SERR + LG EDM sim | SERR + EDM | MARL +LG EDM sim | MARL + EDM |
|---|---|---|---|---|
| 1 | 0 | 0 | 1 | 0 |
| 2 | 73 | 20 | 0 | 0 |
| 3 | 0 | 0 | 0 | 0 |
| 4 | 0 | 0 | 0 | 0 |
| 5 | 0 | 40 | 0 | 80 |
| 6 | 0 | 0 | 99 | 20 |
| 7 | 0 | 0 | 0 | 0 |
| 8 | 22 | 0 | 0 | 0 |
| 9 | 0 | 0 | 0 | 0 |
| 10 | 0 | 0 | 0 | 0 |
| 11 | 0 | 0 | 0 | 0 |
| 12 | 0 | 20 | 0 | 0 |
| 13 | 0 | 0 | 0 | 0 |
| 14 | 0 | 0 | 0 | 0 |
| 15 | 5 | 20 | 0 | 0 |
Additional files
-
Supplementary file 1
Literature review data with published spiralian topologies.
- https://cdn.elifesciences.org/articles/110607/elife-110607-supp1-v1.xlsx
-
MDAR checklist
- https://cdn.elifesciences.org/articles/110607/elife-110607-mdarchecklist1-v1.docx