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. 2022 Aug 24:13:909768.
doi: 10.3389/fpls.2022.909768. eCollection 2022.

An open and continuously updated fern tree of life

Affiliations

Affiliations

  • 1 Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Tokyo, Japan.
  • 2 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC, United States.
  • 3 Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City, Mexico.
  • 4 Department of Integrated Biosciences, Graduate School of Frontier Sciences, The University of Tokyo, Chiba, Japan.
  • 5 Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Chiba, Japan.
  • 6 Atmosphere and Ocean Research Institute, The University of Tokyo, Chiba, Japan.
  • 7 Institute for Quantitative Biosciences, The University of Tokyo, Tokyo, Japan.
  • 8 Collaborative Research Institute for Innovative Microbiology, The University of Tokyo, Tokyo, Japan.

An open and continuously updated fern tree of life

Joel H Nitta et al. Front Plant Sci. .
. 2022 Aug 24:13:909768.
doi: 10.3389/fpls.2022.909768. eCollection 2022.

Affiliations

  • 1 Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Tokyo, Japan.
  • 2 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC, United States.
  • 3 Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City, Mexico.
  • 4 Department of Integrated Biosciences, Graduate School of Frontier Sciences, The University of Tokyo, Chiba, Japan.
  • 5 Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Chiba, Japan.
  • 6 Atmosphere and Ocean Research Institute, The University of Tokyo, Chiba, Japan.
  • 7 Institute for Quantitative Biosciences, The University of Tokyo, Tokyo, Japan.
  • 8 Collaborative Research Institute for Innovative Microbiology, The University of Tokyo, Tokyo, Japan.

Abstract

Ferns, with about 12,000 species, are the second most diverse lineage of vascular plants after angiosperms. They have been the subject of numerous molecular phylogenetic studies, resulting in the publication of trees for every major clade and DNA sequences from nearly half of all species. Global fern phylogenies have been published periodically, but as molecular systematics research continues at a rapid pace, these become quickly outdated. Here, we develop a mostly automated, reproducible, open pipeline to generate a continuously updated fern tree of life (FTOL) from DNA sequence data available in GenBank. Our tailored sampling strategy combines whole plastomes (few taxa, many loci) with commonly sequenced plastid regions (many taxa, few loci) to obtain a global, species-level fern phylogeny with high resolution along the backbone and maximal sampling across the tips. We use a curated reference taxonomy to resolve synonyms in general compliance with the community-driven Pteridophyte Phylogeny Group I classification. The current FTOL includes 5,582 species, an increase of ca. 40% relative to the most recently published global fern phylogeny. Using an updated and expanded list of 51 fern fossil constraints, we find estimated ages for most families and deeper clades to be considerably older than earlier studies. FTOL and its accompanying datasets, including the fossil list and taxonomic database, will be updated on a regular basis and are available via a web portal (https://fernphy.github.io) and R packages, enabling immediate access to the most up-to-date, comprehensively sampled fern phylogeny. FTOL will be useful for anyone studying this important group of plants over a wide range of taxonomic scales, from smaller clades to the entire tree. We anticipate FTOL will be particularly relevant for macroecological studies at regional to global scales and will inform future taxonomic systems with the most recent hypothesis of fern phylogeny.

Keywords: PPGI; fern; phylogeny; plastome; pteridophyte; rbcL.

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Conflict of interest statement

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Figures

FIGURE 1

FIGURE 1

Number of fern species in…

FIGURE 1

Number of fern species in GenBank by year and genomic compartment, 1990–2021. Points…

FIGURE 1
Number of fern species in GenBank by year and genomic compartment, 1990–2021. Points indicate number of species sampled in selected studies of global fern phylogeny (Hasebe et al., 1995; Schuettpelz and Pryer, 2007; Lehtonen, 2011; Testo and Sundue, 2016; this study). This study plotted as 2021 but includes data until 2022-04-15. Schuettpelz and Pryer (2007) did not attempt exhaustive sampling but rather proportional sampling according to lineage size. The relatively small increase in number of species in 2021 may be due to accessions that were still embargoed at the time of writing. Taxonomy of GenBank species follows NCBI (Federhen, 2012). Only accessions identified to species included; environmental samples, hybrid formulas, and names with “aff.” or “cf.” annotations excluded.
FIGURE 2

FIGURE 2

Summary of workflow to construct…

FIGURE 2

Summary of workflow to construct the fern tree of life (FTOL). The workflow…

FIGURE 2
Summary of workflow to construct the fern tree of life (FTOL). The workflow is automated except for steps in boxes with red outlines and a hand symbol. Numbers of sequences and species are approximate. Sequences originating from whole plastomes are in blue; sequences typically obtained by Sanger sequencing are in orange. For details of each step, see Materials and methods.
FIGURE 3

FIGURE 3

Fern tree of life (FTOL).…

FIGURE 3

Fern tree of life (FTOL). Tree rooted with algae. Inset plot shows coverage…

FIGURE 3
Fern tree of life (FTOL). Tree rooted with algae. Inset plot shows coverage by major clade (order or suborder). Bold part of each clade name is its code, which is also indicated on the tree. Numbers next to each bar show sampled species out of total number of species. Taxonomy follows Pteridophyte Phylogeny Group I (2016).
FIGURE 4

FIGURE 4

Fern tree of life (FTOL)…

FIGURE 4

Fern tree of life (FTOL) backbone phylogeny. One exemplar tip is shown per…

FIGURE 4
Fern tree of life (FTOL) backbone phylogeny. One exemplar tip is shown per family (all families were found to be monophyletic; see Results). Ultrafast bootstrap support values (%) shown at nodes; unlabeled nodes are 100%. Outgroup (seed plants, lycophytes, bryophytes, and algae) not shown. Colors of major clades (orders or suborders) correspond to those used in Figure 3. Taxonomy follows Pteridophyte Phylogeny Group I (2016); informal clade names in lowercase.
FIGURE 5

FIGURE 5

Stem age of fern families…

FIGURE 5

Stem age of fern families (Ma) estimated by selected studies. For studies that…

FIGURE 5
Stem age of fern families (Ma) estimated by selected studies. For studies that used methods with confidence intervals, error bars indicate lower and upper 95% highest posterior density levels and point indicates median (Rothfels et al., 2015; Testo and Sundue, 2016). For other studies, point indicates best (most likely) estimate (Schuettpelz and Pryer, 2009; this study). Codes in parentheses after family names indicate major clade as in Figure 3. Period name abbreviations as follows: O (Ordovician), S (Silurian), D (Devonian), C (Carboniferous), P (Permian), Tr (Triassic), J (Jurassic), K (Cretaceous), Pg (Paleogene), Ng (Neogene).

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