Multi-locus phylogeny of Bryoria reveals recent diversification and unexpected diversity in section Divaricatae
Data files
Apr 29, 2026 version files 386.54 KB
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Bryoria_its_mcm7_igs_gapdh_mtssu_tsr1_align.fas
266.42 KB
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GAPDH_Divaricatae.fas
28.57 KB
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igs_Divaricatae.fas
14.06 KB
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its_Divaricatae.fas
22.93 KB
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mcm7_Divaricatae.fas
19.80 KB
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mtssu_Divaricatae.fas
21.02 KB
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README.md
3.67 KB
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Tsr1_Divaricatae.fas
10.07 KB
Abstract
We used information on six gene regions - three nuclear protein-coding markers (Mcm7, GAPDH and Tsr1), two nuclear ribosomal markers (ITS rDNA and IGS rDNA), and a partial mitochondrial small subunit (mtSSU rDNA) - to examine infrageneric relationships in the genus Bryoria (Parmeliaceae, Lecanoromycetes) and to assess species delimitation in Bryoria bicolor/B. tenuis group in section Divaricatae. Our phylogenetic analyses based on the combined data set of six gene loci support the monophyly of sections Americanae, Divaricatae and Implexae, while section Bryoria is polyphyletic and groups in two clades. Species from Bryoria clade 1 are placed in an emended section Americanae. Our study reveals that section Divaricatae is young (c. 5 My) and undergoing diversification especially in South East Asia and western North America. Separate phylogenetic analyses of section Divaricatae using ITS produced a topology congruent with the current species concept. However, the remaining gene regions produced poorly resolved phylogenetic trees and the different species delmitation methods also gnerated highly inconsistent resluts, congruent with other studies that highlight the difficulty of species delmitation in groups with recent and rapid radiation. Based on our results we describe the new species B. ahtiana sp. nov., characterized by its bicolorous, caespitose, widely divergent thallus, conspicuously thickening main stems, well developed secondary branches, and rather sparse third-order branchlets. Another new lineage, here referred to as B. tenuis s. lat., is restricted to western North America and may represent a new species recently diverged from B. tenuis s. str., though further work is needed.
Description of the data and file structure
This submission includes seven datasets:
Bryoria_its_mcm7_igs_gapdh_mtssu_tsr1_align.fas. This is a six-locus data set to examine infrageneric classification of the lichen-forming fungal genus Bryoria. GAPDH = glyceraldehyde 3-phosphate dehydrogenase gene, IGS = intergenic spacer of the nuclear rDNA, its = nuclear ribosomal internal transcribed spacer region, mcm7 = minichromosome maintenance protein 7 gene, mtSSU = small subunit of the mitochondrial ribosomal DNA, Tsr1 = ribosome biogenesis protein
The following six data sets were used to examine phylogenetic relationships and species delimitation in section Divaricatae.
GAPDH_Divaricatae.fas.
igs_Divaricatae.fas
its_Divaricatae.fas
mcm7_Divaricatae.fas
mtssu_Divaricatae.fas
Tsr1_Divaricatae.fas
These are nucelotide sequences alignments (A=adenine, C=cytosine, G=guanine, T=thymine, N=ambiguous, -=deletion).
Sharing/Access information
Links to other publicly accessible locations of the data:
- Sequences are deposited in GenBank (https://www.ncbi.nlm.nih.gov/genbank/)
Data was derived from the following sources:
- Already published sequences were obtained from GenBank (https://www.ncbi.nlm.nih.gov/genbank/). New sequences were obtained from own material and herbarium specimens deposited in H, UPS, UBC, KPABG, OULU and KUN (https://sweetgum.nybg.org/science/ih/)
Code/Software
For all seven data sets, we performed maximum parsimony, maximum likelihood, and, Bayesian analyses. Parsimony analyses were performed in TNT version 1.1 for Windows (Goloboff et al. 2008) using the option Traditional Search with the following settings: random addition of sequences with 100 replicates and TBR branch swapping algorithm. Ten trees were saved for each replicate. The bootstrapping method as implemented in TNT was used with 1000 replicates to estimate node support. Maximum likelihood analyses were performed with RAxML v.8.1.15 (Stamatakis 2014) located at CSC IT center for Science (https://www.csc.fi/). We divided the data set into 23 partitions: ITS1, 5.8S, ITS2, IGS, mtSSU, each three codon positions of Mcm7, GAPDH and Tsr1, and introns of GAPDH. These partitions were analysed under the universal GTR-GAMMA model. Node support was estimated with 1000 bootstrap replicates using the rapid bootstrap algorithm.
For the Bayesian analyses, the optimal substitution model for each locus was calculated in jmodeltest (Posada 2008), using Akaike information criterion (AIC). The models selected were: TrNef+G for ITS1, IGS rDNA, GAPDH; JC for 5.8S; K80+G for ITS2; TrNef+I+G for Mcm7; SYM+I+G for mtSSU; SYM+G for Tsr1.
The Bayesian analyses were run in MrBayes 3.2.6 (Ronquist et al. 2012) in CIPRES Science Gateway v. 3.1 (Miller et al. 2010). For the concatenated analysis 23 partitions were considered and the models selected by jmodeltest were used. The posterior probabilities were approximated by sampling trees using Markov Chain Monte Carlo (MCMC). Two simultaneous runs with 20 000 000 generations each, starting with a random tree and employing 4 simultaneous chains, were executed. Every 1 000th tree was saved into a file. The first 25% trees were deleted as burn-in. The convergence between chains was assessed in Tracer v. 1.7 (Rambaut et al. 2018), plotting the likelihood versus generation number and the average standard deviation of split frequencies ( 0.01).
A total of 71 Bryoria specimens from North America, Europe and Asia were used in the molecular phylogenies. Six markers were used: ITS rDNA regions, IGS rDNA, partial mtSSU rDNA gene, partial Mcm7 gene, partial GAPDH gene and partial Tsr1 gene. Seven data sets were prepared: 1) six-locus data set to examine infrageneric classification. The data set included 64 ingroup specimens and Usnea dasopoga was used as an outgroup taxon; 2-7) separate ITS (42 specimens), IGS (30 specimens), mtSSU (30 specimens), GAPDH (27 specimens), Mcm7 (31 specimens) and Tsr1 (15 specmiens) data sets to examine phylogeny and species delimitation within the Bryoria bicolor/B. tenuis group in section Divaricate, Bryoria americana was used as an outgroup taxon based on the phylogenies by Myllys et al. (2011, 2016). Gene regions were aligned separately with MUSCLE v.3.8.31 (Edgar 2004) using EMBL-EBI's freely available web service (http://www.ebi.ac.uk/Tools/msa/muscle/). Maximum parsimony, maximum likelihood, and, Bayesian analyses were performed for all seven data sets.
PhyDE - Phylogenetic Data Editor: http://www.phyde.de/docu/docu.html
