JSE2016

Journal of Systematics
JSE and Evolution doi: 10.1111/jse.12229
Research Article
A community-derived classification for extant lycophytes

and ferns
The Pteridophyte Phylogeny Group
Recommended citation: PPG I (2016). This project was organized by Eric Schuettpelz1*,
Harald Schneider2*, Alan R. Smith3, Peter Hovenkamp4, Jefferson Prado5, Germinal Rouhan6,
Alexandre Salino7, Michael Sundue8, Thaıs Elias Almeida9, Barbara Parris10, Emily B. Sessa11, Ashley R. Field12,
Andre Luıs de Gasper13, Carl J. Rothfels14, Michael D. Windham15, Marcus Lehnert16, Benjamin Dauphin17,
Atsushi Ebihara18, Samuli Lehtonen19, Pedro Bond Schwartsburd20, Jordan Metzgar21, Li-Bing Zhang22,
Li-Yaung Kuo23, Patrick J. Brownsey24, Masahiro Kato18, and Marcelo Daniel Arana25; with additional
contributions from (in alphabetical order): Francine C. Assis7, Michael S. Barker26, David S. Barrington8,
Ho-Ming Chang27, Yi-Han Chang28, Yi-Shan Chao29, Cheng-Wei Chen30, De-Kui Chen31, Wen-Liang Chiou32,
Vinıcius Antonio de Oliveira Dittrich33, Yi-Fan Duan34, Jean-Yves Dubuisson35, Donald R. Farrar36, Susan Fawcett8,
Jose Marıa Gabriel y Galan37, Luiz Armando de Ara es-Neto7, Jason R. Grant17, Amanda L. Grusz38,
ujo Go
Christopher Haufler39, Warren Hauk40, Hai He31, Sabine Hennequin35, Regina Yoshie Hirai5, Layne Huiet15,
n44, Chun-Xiang Li45, Fay-Wei Li15,
Michael Kessler41, Petra Korall42, Paulo H. Labiak43, Anders Larsson42, Blanca Leo
Melanie Link-Pe rez , Hong-Mei Liu , Ngan Thi Lu , Esteban I. Meza-Torres , Xin-Yuan Miao45, Robbin Moran50,
46 47 48 49
Claudine Massi Mynssen51, Nathalie Nagalingum52, Benjamin Øllgaard53, Alison M. Paul54, Jovani B. de S. Pereira55,
Leon R. Perrie24, Mo nica Ponce56, Tom A. Ranker57, Christian Schulz55, Wataru Shinohara58, Alexander Shmakov59,
Erin M. Sigel , Filipe Soares de Souza7, Lana da Silva Sylvestre60, Weston Testo8, Luz Amparo Triana-Moreno61,
1
Chie Tsutsumi18, Hanna Tuomisto19, Ivan A. Valdespino62, Alejandra Vasco63, Raquel Stauffer Viveros7,
Alan Weakley64, Ran Wei65, Stina Weststrand42, Paul G. Wolf66, George Yatskievych44, Xiao-Gang Xu34,
Yue-Hong Yan67, Liang Zhang68, Xian-Chun Zhang65, and Xin-Mao Zhou22.
1
Department of Botany, National Museum of Natural History, Smithsonian Institution
2
Institute of Ecology and Evolution, School of Life Sciences, Sun Yat-Sen University & Department of Life Sciences, Natural History Museum,
London
3
University Herbarium, University of California, Berkeley
4
Research & Education, Naturalis Biodiversity Center, Leiden
5
Herbario SP, Instituto de Botânica, S~ao Paulo
6
Institut de Syst
ematique, Evolution, Biodiversit e, Sorbonne Universit es, Mus
eum National d’Histoire Naturelle, Herbier National, Paris
7
Departamento de Botânica, Universidade Federal de Minas Gerais
8
Pringle Herbarium, Department of Plant Biology, University of Vermont
9
Herbario HSTM, Universidade Federal do Oeste do Par a
10
Fern Research Foundation, Kerikeri
11
Department of Biology, University of Florida
12
Queensland Herbarium, Department of Science and Innovation and Australian Tropical Herbarium, James Cook University
13
Departamento de Ci^ encias Naturais, Universidade Regional de Blumenau
14
University Herbarium and Department of Integrative Biology, University of California, Berkeley
15
Department of Biology, Duke University
16
Nees-Institut f€
ur Biodiversit€
at der Pflanzen, Rheinische Friedrich-Wilhelms-Universit€at Bonn
17
Laboratoire de Botanique Evolutive, Institut de Biologie, Universit
e de Neuchâtel
18
Department of Botany, National Museum of Nature and Science, Tokyo
19
Department of Biology, University of Turku
20
Departamento de Biologia Vegetal, Universidade Federal de ViSc osa
21
Museum of the North, University of Alaska Fairbanks
22
Missouri Botanical Garden
23
Institute of Ecology and Evolutionary Biology, National Taiwan University
24
Collections, Research, and Learning, Museum of New Zealand Te Papa Tongarewa
25
Departamento de Ciencias Naturales, Universidad Nacional de Rıo Cuarto
November 2016 | Volume 54 | Issue 6 | 563–603 © 2016 Institute of Botany, Chinese Academy of Sciences
564 The Pteridophyte Phylogeny Group
26
Department of Ecology and Evolutionary Biology, University of Arizona
27
Division of Botany, Taiwan Endemic Species Research Institute
28
Hengchun Research Center, Taiwan Forestry Research Institute
29
Department of Biomedical Science and Environmental Biology, Kaohsiung Medical University
30
Division of Botanical Garden, Taiwan Forestry Research Institute
31
College of Life Sciences, Chongqing Normal University
32
Herbarium, Taiwan Forestry Research Institute
33
Departamento de Bot^ anica, Universidade Federal de Juiz de Fora
34
Department of Botany, College of Biology and the Environment, Nanjing Forestry University
35
Institut de Syst
ematique, Evolution, Biodiversit e, Sorbonne Universit
es, Universit
e Pierre et Marie Curie, Paris
36
Department of Ecology, Evolution and Organismal Biology, Iowa State University
37
Departamento de Biologıa Vegetal I, Universidad Complutense de Madrid
38
Department of Biology, University of Minnesota Duluth
39
Department of Ecology and Evolutionary Biology, University of Kansas
40
Department of Biology, Denison University
41
Department of Systematic and Evolutionary Botany, University of Zurich
42
Department of Organismal Biology, Evolutionary Biology Centre, Uppsala University
43
Departamento de Bot^ anica, Universidade Federal do Paran a
44
Plant Resources Center, University of Texas at Austin
45
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences
46
Department of Botany and Plant Pathology, Oregon State University
47
Shenzhen Key Laboratory of Southern Subtropical Plant Diversity, Fairylake Botanical Garden, Chinese Academy of Sciences
48
Department of Botany, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, Hanoi
49
Instituto de Botanica del Nordeste, Universidad Nacional del Nordeste, Consejo Nacional de Investigaciones Cientıficas y T
ecnicas
50
New York Botanical Garden
51
Diretoria de Pesquisa, Instituto de Pesquisa Jardim Bot^ anico do Rio de Janeiro
52
National Herbarium of New South Wales, Royal Botanic Gardens and Domain Trust
53
Section for Ecoinformatics and Biodiversity, Aarhus University
54
Department of Life Sciences, Natural History Museum, London
55
Department of Evolution and Biodiversity of Plants, Ruhr-Universit€ at Bochum
56
Instituto de Botanica Darwinion, Consejo Nacional de Investigaciones Cientıficas y T ecnicas, Buenos Aires
57
Department of Botany, University of Hawaii at Manoa
58
Faculty of Education, Kagawa University
59
South-Siberian Botanical Garden, Altai State University
60
Departamento de Botânica, Universidade Federal do Rio de Janeiro
61
Departamento de Ciencias Biol ogicas, Universidad de Caldas
62
Departamento de Botanica, Universidad de Panam a
63
Instituto de Biologıa, Universidad Nacional Aut onoma de M exico
64
UNC Herbarium, North Carolina Botanical Garden, University of North Carolina at Chapel Hill
65
Institute of Botany, Chinese Academy of Sciences
66
Department of Biology, Utah State University
67
Shanghai Chenshan Botanical Garden, Shanghai Chenshan Plant Science Research Center, Chinese Academy of Sciences
68
Kunming Institute of Botany, Chinese Academy of Sciences
*Authors for correspondence. E-mail: schuettpelze@si.edu and haralds@mail.sysu.edu.cn
Received 7 October 2016; Accepted 1 November 2016; Article first published online 13 December 2016
Abstract Phylogeny has long informed pteridophyte classification. As our ability to infer evolutionary trees has
improved, classifications aimed at recognizing natural groups have become increasingly predictive and stable. Here,
we provide a modern, comprehensive classification for lycophytes and ferns, down to the genus level, utilizing a
community-based approach. We use monophyly as the primary criterion for the recognition of taxa, but also aim to
preserve existing taxa and circumscriptions that are both widely accepted and consistent with our understanding of
pteridophyte phylogeny. In total, this classification treats an estimated 11 916 species in 337 genera, 51 families, 14
orders, and two classes. This classification is not intended as the final word on lycophyte and fern taxonomy, but
rather a summary statement of current hypotheses, derived from the best available data and shaped by those most
familiar with the plants in question. We hope that it will serve as a resource for those wanting references to the
recent literature on pteridophyte phylogeny and classification, a framework for guiding future investigations, and a
stimulus to further discourse.
Key words: classification, ferns, lycophytes, monophyly, phylogeny, pteridophytes.
J. Syst. Evol. 54 (6): 563–603, 2016 www.jse.ac.cn

PPG I 565
Free-sporing vascular plants comprise two distinct evolution- 2011) and ultimately lead to greater stability. Although we
ary lineages—lycophytes and ferns (e.g., Kenrick & Crane, acknowledge the validity of some arguments presented in
1997; Pryer et al., 2001)—that share a unique life cycle with favor of maintaining paraphyletic taxa (see, e.g., H€ orandl &
independent gametophyte and sporophyte phases (e.g., Stuessy, 2010), we ultimately reject this practice and aim to
Haufler et al., 2016). Classifications of these plants—often recognize only monophyletic lineages at the genus level and
referred to collectively as pteridophytes despite their above. There are many reasons why monophyly might not be
phylogenetic separation—have long been grounded in expected at the species level. This is especially true when
evolutionary principles and have reflected perceived evolu- polyploid speciation is rampant and, among pteridophytes, it
tionary trends (e.g., Copeland, 1947; Pichi Sermolli, 1973, has been estimated that nearly a third of all speciation events
1977b; Kramer & Green, 1990; Smith, 1995; Smith et al., 2006b; are correlated with an increase in ploidy (Wagner & Wagner,
Christenhusz & Chase, 2014). However, these classifications 1980; Wood et al., 2009). We acknowledge the difficulties
were often conflicting, in large part due to a paucity of inherent in translating evolutionary tree structures into
information concerning pteridophyte relationships and a lack hierarchical rank-based classifications. However, because
of consensus regarding patterns of morphological evolution. such classifications have been implemented for centuries and
The increased availability of molecular data and advances in continue to be used extensively, we do not subscribe to the
phylogenetic methods have revolutionized our understanding alternative approach (disruptive, from the standpoint of
of lycophyte and fern relationships. Today, as a consensus stability) recommended in the PhyloCode (Cantino & de
emerges concerning the evolutionary history of these Queiroz, 2000). Due to the many challenges of incorporating
lineages, classifications aimed at recognizing their natural fossil taxa, this first installment of the PPG classification focuses
groups are becoming increasingly “predictive” (Stuessy, exclusively on extant lycophytes and ferns. Although some
2009) and, we hope, stable. fossils could easily be accommodated, the phylogenetic
The ordinal and familial scheme by Smith et al. (2006b) was affinities of most extinct plants are rather unclear. Many fossil
the first higher-level pteridophyte classification published in taxa represent distinct evolutionary lineages and their inclusion
the molecular era. Although this work focused exclusively on in the PPG I classification would not only require revised
ferns, these plants—also referred to as “monilophytes” in circumscriptions, but almost certainly the recognition of new
some publications (e.g., Pryer et al., 2004; Smith et al., 2006b; families, orders, and even classes.
Schneider et al., 2009; here we simply use “ferns”)—account In general, the PPG I classification seeks to preserve existing
for nearly 90% of extant pteridophyte diversity. Founded on taxa and circumscriptions that are widely accepted and
the principle of monophyly, while also recognizing the consistent with our understanding of lycophyte and fern
importance of maintaining well-established names and phylogeny. Although monophyly is our primary criterion for
circumscriptions, the Smith et al. (2006b) classification the recognition of taxa (Backlund & Bremer, 1998), we adopt a
established a new standard in fern taxonomy. Over the past conservative approach that minimizes name changes in cases
decade, many important advances have been made in our where data are insufficient to circumscribe what we judge to
understanding of relationships, some of which are reflected in be stable monophyletic groups; this is with the understanding
subsequent higher-level schemes (e.g., Christenhusz et al., that some currently accepted taxa may ultimately be revealed
2011; Rothfels et al., 2012b). A recent scheme by Christenhusz to be non-monophyletic. In making decisions, we secondarily
& Chase (2014) was also consistent with an understanding of weigh morphological diagnosability and homogeneity, and
fern phylogeny but represented a considerable departure in hierarchical equivalency in terms of both age and diversity.
terms of stability and has not been widely adopted. This classification is not meant to be the final word on
Here, we provide a modern, comprehensive classification pteridophyte taxonomy, but rather a summary statement of
for lycophytes and ferns, following the main tenets of the current hypotheses, derived from the best available data and
Smith et al. (2006b) work but utilizing a democratic shaped by those most familiar with the plants in question. We
community-based approach. To accomplish this goal, we hope that it will serve as a resource for those wanting
have established the Pteridophyte Phylogeny Group (PPG), references to recent literature on lycophyte and fern
based loosely on the model employed for flowering plant phylogeny, a framework for guiding future investigations,
classification (APG, 1998; APG II, 2003; APG III, 2009; APG IV, and a stimulus to further discourse. Although we welcome its
2016). Below, we outline our general philosophy and use by systematists, other scientists, herbaria, governmental
approach, summarize our outcomes, and present a consensus agencies, and others with a professional or non-professional
classification for all pteridophytes, from the rank of class to interest in plants, we recognize that a single system may not
that of genus. serve all users and we are strongly opposed to this
classification, or any other classification, being imposed on
investigators, authors, editors, or reviewers. Disagreements
exist even among the contributors to this PPG I classification,
Philosophy as noted below.
Biological classifications are fundamentally tools for communi-
cation about biodiversity (e.g., Stuessy, 2009). The stability of
names is thus of great importance, and it is critical to consider
existing classifications when determining taxa worthy of
Approach
recognition and the ranks at which to treat them. A focus on Establishment of the Pteridophyte Phylogeny Group (PPG)
natural groups is similarly important, as it results in classi- began through promotion of the concept at international
fications that reflect evolutionary history (Schmidt-Lebuhn, conferences (including the 2015 Next Generation Pteridology
www.jse.ac.cn J. Syst. Evol. 54 (6): 563–603, 2016

conference) and posts to society websites and e-mail lists, of the taxon; (4) a brief statement concerning its monophyly;
including those of the American Fern Society (AFS) and the (5) a list of any included taxa (i.e., synonyms); (6) the number
International Association of Pteridologists (IAP). After a series of species attributed to the taxon; and (7) any additional
of informal discussions among the early respondents, an comments.
electronic mailing list was established to enhance community The classification presented herein reflects our current
engagement and facilitate discussion. As the project gained understanding of lycophyte and fern phylogeny (see refer-
momentum, the community of collaborators expanded; ences below). But, ultimately, circumscriptions adopted
currently, the PPG is a global community of 94 pteridologists. herein are all subject to reconsideration, given new evidence.
The classification presented herein is community-derived, Future research will enable greater insight into pteridophyte
and the taxa recognized have the support of a majority (or a evolution and our classification will need to accommodate
plurality, in select instances) of the contributors. Decisions more robust hypotheses. To this end, we envision formal
were made using an inclusive, transparent, and straightfor- updates to the PPG I classification, with interim improvements
ward approach that typically involved: (1) an initial conversa- tracked on a PPG website. Such adjustments (most likely a
tion among members of a focused subcommittee consisting series of small incremental changes) will be carried out via the
of experts willing to contribute to the discussion; (2) process outlined above, with initial proposals generated by
formulation of a draft proposal by the subcommittee; (3) smaller groups that are subsequently discussed and approved
discussion of the draft proposal among all members of the or rejected by the broader PPG.
PPG community; (4) production of a revised proposal; and (5)
approval of the revised proposal through a formal vote or
simple acclamation.
For the highest-level taxa, a dialogue was initiated across
Outcomes
the whole of the PPG. After discussion of the advantages and We recognize two pteridophyte classes: Lycopodiopsida
disadvantages of various options, one system (recognizing (lycophytes) and Polypodiopsida (ferns). These are distinct
classes and subclasses) emerged as the clear preference. A lineages within the tracheophyte tree of life, with ferns
vote was taken to ensure there was majority support. A similar resolved as more closely related to seed plants than to
approach was used later to reach decisions concerning the lycophytes (Fig. 1; Kenrick & Crane, 1997; Pryer et al., 2001).
informal names to be applied to major clades. Within Lycopodiopsida, we further recognize three orders
To determine the orders, suborders, and families to be (Lycopodiales, Isoëtales, and Selaginellales). Order Lycopo-
recognized, an organizing subcommittee developed the initial diales includes one family and 16 genera, whereas orders
proposal. This was then shared electronically with all Isoëtales and Selaginellales each contain a single monogeneric
contributors, along with a survey developed to gauge family.
acceptability and highlight areas needing further discussion. Within Polypodiopsida, we recognize four subclasses:
With the results of this survey in hand, discussion proceeded Equisetidae (horsetails); Ophioglossidae; Marattiidae; and
among all contributors. Following this dialogue, a revised poll Polypodiidae (leptosporangiates). Extant Equisetidae includes
was constructed that presented alternatives in areas of a single order, a single small family, and a single genus
disagreement. All orders, suborders, and families recognized (Equisetum L.). Subclass Ophioglossidae encompasses two
herein ultimately received majority support, ranging from just orders, each with a single family, and a total of 12 genera.
over 50% to unanimous approval. Marattiidae includes just one order, one family, and six
After settling on a list of higher-level taxa to be recognized, genera.
members of the PPG were asked (via an online survey) Subclass Polypodiidae comprises the vast majority of extant
whether they felt that the generic classification within fern diversity (Fig. 1). Here, we recognize seven orders
particular clades was in need of attention. In cases where a (Osmundales, Hymenophyllales, Gleicheniales, Schizaeales,
unanimous consensus existed, the organizers implemented Salviniales, Cyatheales, and Polypodiales), with the Polypodiales
the prevailing generic classification. If a clade were identified subsequently divided into six suborders (Saccolomatineae,
as needing attention, a subcommittee was established, Lindsaeineae, Pteridineae, Dennstaedtiineae, Aspleniineae, and
consisting of contributors with experience in the particular Polypodiineae). Within Osmundales, we recognize a single small
lineage. family with six genera. Order Hymenophyllales also includes just
Clade-focused subcommittees, under the leadership of one family, but it is considerably larger and encompasses nine
subcommittee heads (selected by subcommittee members), genera. Gleicheniales and Schizaeales each constitute three
discussed alternative generic classifications, reached a relatively small families, with ten and four total genera,
consensus, and presented a proposal for approval by the respectively. The order Salviniales comprises two families and
whole of the PPG. Any concerns were mediated via open five genera, and the order Cyatheales encompasses seven small
dialogue through the PPG e-mail list. families, with a total of ten genera, plus the larger Cyatheaceae
After arriving at a system of classes, subclasses, orders, with three genera.
suborders, families, and genera to be recognized, subcommit- Within the Polypodiales, suborder Saccolomatineae in-
tee heads (with the assistance of other contributors) were cludes a single small monogeneric family (Saccolomataceae).
then tasked with compiling the following pertinent informa- Suborder Lindsaeineae comprises two monogeneric families
tion for each included taxon: (1) the author of the name and plus the larger Lindsaeaceae with seven genera. We recognize
the protologue reference; (2) for genera, the type species and a single large family within Pteridineae, which includes 52
basionym, along with the lectotypification reference (where genera. Suborder Dennstaedtiineae also includes a single
applicable); (3) a brief statement outlining the circumscription family, with ten genera. The remaining two suborders,

PPG I 567
Fig. 1. Summary tracheophyte phylogeny, depicting relationships among lycophyte and fern families recognized in this PPG I
classification. Composite topology is derived from the results of numerous phylogenetic studies (e.g., Pryer et al., 2001, 2004; Korall et al.,
2006; Schuettpelz & Pryer, 2007; Rai & Graham, 2010; Lehtonen, 2011; Rothfels et al., 2012a, 2015; Knie et al., 2015; Zhang & Zhang, 2015).
Most nodes have received consistently strong support; dotted lines indicate areas of considerable uncertainty. For each family, we note
the total number of genera recognized in PPG I and the sum of species estimates for these genera (terminal clade height is roughly
proportional to diversity for families with more than 100 species). Where applicable, informal clade names are provided in parentheses.

Aspleniineae (eupolypods II) and Polypodiineae (eupolypods most analyses of morphological data (Kenrick & Crane, 1997;
I), are exceedingly diverse, together accounting for well over Schneider et al., 2009).
half of extant pteridophyte diversity. Suborder Aspleniineae Pteridophyte classification has been exceedingly stable at
encompasses four large families, with a total of 59 genera, and the ordinal level over the past decade. Smith et al. (2006b)
seven smaller families, with 14 genera and fewer than 100 recognized 11 orders within ferns, one for each of four major
species. Finally, suborder Polypodiineae comprises three large eusporangiate lineages and seven within leptosporangiates.
families, with a total of 98 genera, and six small families, with Christenhusz et al. (2011) and Christenhusz & Chase (2014)
ten genera. adopted these same 11 fern orders plus three orders for
In all, this PPG I classification treats an estimated 11 916 lycophytes (not treated by Smith et al., 2006b). At the ordinal
species in 337 genera, 51 families, 14 orders, and 2 classes. level, the PPG I classification does not differ at all from the
In some cases, additional divisions are made to recognize earlier classifications, but we do introduce a new informal
subclasses and suborders (as noted above), as well as name for one large order. Polypodiales, which accounts for
subfamilies (see classification below). Classes, subclasses, more than 80% of fern diversity, has previously been referred
orders, suborders, and families, are organized phylogenet- to as “polypods” (Schneider et al., 2004b; Smith et al., 2006b).
ically, generally with increasing species diversity (Fig. 1). However, “polypods” has also been used to refer to the
Within each family, subfamilies, if recognized, are orga- members of the family Polypodiaceae that do not belong to
nized by increasing species diversity; genera are then the distinct (and diverse) “grammitid” clade (Smith, 1995;
organized alphabetically within each family or subfamily, as Sundue et al., 2014). Here, we aim to avoid confusion by using
applicable. a more descriptive informal name for the more inclusive clade
(i.e., that equivalent to Polypodiales). We refer to this clade as
“cathetogyrates” (a term first used by Bernhardi, 1806), in
reference to the unique sporangium—with a vertical ring of
Discussion annular cells—that characterizes this lineage. Within the
As noted above, the PPG I classification presented here Polypodiales, we recognize six suborders, providing formal
reflects our current, collective understanding of lycophyte names for “eupolypods I”, “eupolypods II”, and other clades.
and fern phylogeny. It takes into consideration all studies As this rank has not typically been utilized within ferns, five of
examining the relationships of these plants published through these suborders are new.
mid-2016. The PPG I classification also aims to maintain, where Outside of the Polypodiales, the 25 families recognized in
possible, continuity in the naming and circumscription of this PPG I classification are identical to those in Smith et al.
families and genera as recognized by most pteridologists prior (2006b, for ferns) and Christenhusz et al. (2011). Within the
to the molecular revolution (e.g., Kramer & Green, 1990). In Polypodiales, we recognize 26 families, and there are some
this respect, this study continues the approach employed in stark differences relative to Smith et al. (2006b) and
most recent classifications (e.g., Smith et al., 2006b; Christenhusz et al. (2011). The Lindsaeaceae, Woodsiaceae,
Christenhusz et al., 2011; Rothfels et al., 2012b) but differs Dryopteridaceae, and Lomariopsidaceae sensu Smith et al.
from classifications that favored a reduction in the number of (2006b) have all been found to be paraphyletic in more recent
taxa recognized (e.g., Christenhusz & Chase, 2014). phylogenetic analyses and are here divided into three, seven,
At the highest taxonomic ranks, the PPG I classification is three, and two families, respectively. The present PPG I
wholly compatible with a scheme recently proposed for all of classification additionally recognizes subfamilies within some
life. Ruggiero et al. (2015) treated lycophytes and ferns as of the larger families of lycophytes and ferns, especially those
distinct subphyla (Lycopodiophytina and Polypodiophytina), with a history of subdivision. This has the advantage of
each with a single class (Lycopodiopsida and Polypodiopsida). providing formal names for some clades (e.g., vittarioids and
The present classification recognizes the two classes but does grammitids) that were long recognized as distinct families but
not make recommendations above this rank. Further, the PPG were later subsumed into other families (within which they
I classification agrees with Ruggiero et al. (2015) in recognizing appear to be nested).
four subclasses within ferns (Equisetidae, Ophioglossidae, Unlike recent comprehensive treatments of lycophytes and
Marattiidae, and Polypodiidae). This approach differs from ferns (e.g., Smith et al., 2006b; Christenhusz et al., 2011), the
that of Smith et al. (2006b), in which the four major lineages of PPG I classification extends to the genus level. We formally
ferns were recognized as classes, as well as from that of Chase recognize 337 pteridophyte genera, representing a 50%
& Reveal (2009), where embryophytes were treated collec- increase over the 223 genera included in the pre-molecular
tively as a single class, but is consistent with the scheme used system of Kramer & Green (1990). Although this difference is
in the Tree of Life Web Project (Pryer et al., 2009). Although dramatic, the constituent changes have, in fact, accumulated
two of these higher-level classifications (Smith et al., 2006b; incrementally over the past quarter-century. When existing
Chase & Reveal, 2009) do not provide a formal scientific name generic concepts are found to be in conflict with the results of
for ferns, they do both clearly unite the four fern lineages that molecular phylogenetic analyses, there are, using monophyly
we recognize as subclasses (Equisetidae, Ophioglossidae, as a criterion, two paths to resolution: disintegration (i.e., the
Marattiidae, and Polypodiidae). We do not consider our “splitting” of a non-monophyletic genus into multiple
decision to include them together in a single class to be monophyletic genera to maintain a nested genus); or
controversial, as the corresponding clade (i.e., ferns) has been integration (i.e., the “lumping” of a nested genus with the
resolved consistently in analyses of molecular data (e.g., Pryer genus it renders non-monophyletic). Both approaches have an
et al., 2001, 2004; Qiu et al., 2007; Schneider et al., 2009; Ruhfel impact on stability, as the former results in new descriptions
et al., 2014; Wickett et al., 2014; Rothfels et al., 2015) and even (or reinstatements) and the latter in synonymizations.

PPG I 569
Looking again at the marked increase in the number of Ji et al., 2008; Field et al., 2016). Three subfamilies, 16
pteridophyte genera, it is clear that disintegration has genera, and an estimated 388 species.
typically been favored and some lineages have experienced
greater change than others. Instead of recognizing just four Subfamily Lycopodielloideae W.H.Wagner & Beitel ex
genera within the Lycopodiaceae (sensu Øllgaard, 1990), the B.Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription
PPG I classification takes the now-common approach of sensu Øllgaard (2015). Monophyletic (Field et al.,
subdividing the family into 16 genera (Holub, 1964, 1975, 1983, 2016). Included in Lycopodioideae in Field et al. (2016).
1985, 1991; Wagner & Beitel, 1992; Haines, 2003; Øllgaard, Four genera and an estimated 54 species.
2012a, 2012b, 2015; Field & Bostock, 2013; Øllgaard & Windisch,
2014; Field et al., 2016). Likewise, we mostly follow Almeida Lateristachys Holub, Folia Geobot. Phytotax. 18:
et al. (2016) in our treatment of 30 genera in Thelypteridaceae 440. 1983. Type: Lateristachys lateralis (R.Br.)
(versus five in Smith, 1990) and we also incorporate Holub ( Lycopodium laterale R.Br.). Circumscrip-
considerable subdivision recently proposed for the Hymeno- tion sensu Holub (1983). Monophyletic (Field et al.,
phyllaceae (sensu Ebihara et al., 2006), grammitids (sensu 2016). Four species.
Sundue et al., 2014), and Blechnaceae (sensu Gasper et al.,
2016b). In the majority of these studies, the authors were Lycopodiella Holub, Preslia 36: 20, 22. 1964. Type:
largely able to re-establish preexisting (narrower) concepts Lycopodiella inundata (L.) Holub ( Lycopodium
that were both introduced and discarded prior to the inundatum L.). Circumscription sensu Øllgaard &
molecular revolution. Windisch (2014). Monophyletic (Field et al., 2016).
Of course, more inclusive generic concepts have been 15 species.
favored in some cases. We continue to recognize a broad
Asplenium L. (segregating only Hymenasplenium Hayata, sensu Palhinhaea Franco & Vasc., Bol. Soc. Brot., ser. 2,
Schneider et al., 2004a). And, despite recent subdivisions, the 41: 24. 1967. Type: Palhinhaea cernua (L.) Vasc. &
present PPG I classification ultimately recognizes an inclusive Franco ( Lycopodium cernuum L.). Circumscrip-
Davallia Sm. (sensu Tsutsumi et al., 2016) and Woodsia R.Br. tion sensu Holub (1985, 1991), Øllgaard (2012b),
(sensu Shao et al., 2015). Furthermore, the genus Dryopteris and Øllgaard & Windisch (2014). Monophyletic
Adans. is greatly expanded here, relative to its definition at the (Field et al., 2016). Palhinhaea Franco & Vasc. is
end of the 20th century (Kramer et al., 1990), to include proposed to be conserved against Lepidotis P.
segregates such as Acrophorus C.Presl, Dryopsis Holttum & Beauv. (Greuter & Troia, 2014). 25 species.
P.J.Edwards, Nothoperanema (Tagawa) Ching, and Peranema
D.Don (Zhang & Zhang, 2012; Zhang et al., 2012). Notably, Pseudolycopodiella Holub, Folia Geobot. Phytotax.
resolution via integration, and the resulting recognition of 18: 441. 1983. Type: Pseudolycopodiella caroliniana
larger genera, appears to be much more common in flowering (L.) Holub ( Lycopodium carolinianum L.). Circum-
plants (Humphreys & Linder, 2009). scription sensu Holub (1983). Monophyletic (Field
The PPG I classification presented below does not provide et al., 2016). 10 species.
species lists; however, we do present estimates of species
number for each genus. Interestingly, our genus-based Subfamily Lycopodioideae W.H.Wagner & Beitel ex B.
totals (1338 lycophyte species and 10 578 fern species) are Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription
remarkably similar to the family-based sums provided sensu Øllgaard (2015). Monophyletic (Field et al.,
by Christenhusz & Chase (2014; 1300 lycophytes and 10 535 2016). Including Lycopodielloideae in Field et al. (2016).
ferns). Nine genera and an estimated 58 species.
Austrolycopodium Holub, Folia Geobot. Phytotax.

26(1): 90. 1991. Type: Austrolycopodium magellani-
Classification cum (P.Beauv.) Holub ( Lepidotis magellanica P.
Class Lycopodiopsida Bartl., Ord. Nat. Pl.: 14, 19. 1830. Beauv.). Circumscription sensu Holub (1991).
Circumscription sensu Ruggiero et al. (2015). Monophyletic Monophyletic (Field et al., 2016). Eight species.
(Pryer et al., 2001; Kenrick & Crane, 1997; Rai & Graham, 2010;
Wickett et al., 2014). Three orders, three families, 18 genera, Dendrolycopodium A.Haines, Fam. Huperziac. Ly-
and an estimated 1338 species. copodiac. New England: 84. 2003. Type: Dendro-
lycopodium obscurum (L.) A.Haines ( Lycopodium
A. Order Lycopodiales DC. ex Bercht. & J.Presl, Prir. Rostlin: obscurum L.). Circumscription sensu Haines
272. 1820. Circumscription equivalent to family Lycopo- (2003). Monophyletic (Field et al., 2016). Four
diaceae in this classification. Monophyletic (Wikstro €m & species.
Kenrick, 1997, 2000, 2001; Ji et al., 2008; Field et al., 2016).
One family, 16 genera, and an estimated 388 species. Diphasiastrum Holub, Preslia 47(2): 104. 1975. Type:
Diphasiastrum complanatum (L.) Holub ( Lycopo-
1. Family Lycopodiaceae P.Beauv. in Mirb., Hist. Nat. Veg. dium complanatum L.). Circumscription sensu
4: 293. 1802. Circumscription sensu Øllgaard (2015). Holub (1975, 1985). Monophyletic (Field et al.,
Monophyletic (Wikstro€m & Kenrick, 1997, 2000, 2001; 2016). 20 species.

Diphasium C.Presl ex Rothm., Repert. Spec. Nov. Phylloglossum Kunze, Bot. Zeitung (Berlin) 1: 721.
Regni Veg. 54: 64. 1944. Type: Diphasium jussiaei 1843. Type: Phylloglossum drummondii Kunze.
(Desv.) Rothm. ( Lycopodium jussiaei Desv.). Circumscription sensu Field et al. (2016). Monotypic
Circumscription sensu Holub (1985). Monophyletic (Field et al., 2016).
(Field et al., 2016). Five species.
B. Order Isoëtales Prantl, Lehrb. Bot.: 116, 125. 1874.
Lycopodiastrum Holub ex R.D.Dixit, J. Bombay Nat. Circumscription equivalent to family Isoëtaceae in this
Hist. Soc. 77(3): 540. 1981. Type: Lycopodiastrum classification. Monophyletic (Rydin & Wikstr€ om, 2002;
casuarinoides (Spring) Holub ex R.D.Dixit ( n & Rydin, 2016). One family, one
Hoot et al., 2006; Larse
Lycopodium casuarinoides Spring). Circumscription genus, and about 250 species.
sensu Holub (1983) and Zhang & Iwatsuki (2013).
Monotypic (Field et al., 2016). 2. Family Isoëtaceae Dumort., Anal. Fam. Pl.: 67. 1829.
Circumscription sensu Jermy (1990a). Monophyletic
Lycopodium L., Sp. Pl. 2: 1100. 1753. Lectotype (Rydin & Wikstro€m, 2002; Hoot et al., 2006; Larse
n &
(designated by W.J.Rob., Bull. Torrey Bot. Club 41: Rydin, 2016). One genus and about 250 species.
51. 1914): Lycopodium clavatum L. Circumscription
sensu Haines (2003). Monophyletic (Field et al., Isoëtes L., Sp. Pl. 2: 1100. 1753. Type: Isoëtes lacustris
2016). 15 species. L. Circumscription sensu Jermy (1990a). Monophy-
n & Rydin, 2016). About 250 species.
letic (Larse
Pseudodiphasium Holub, Folia Geobot. Phytotax.
18: 440. 1983. Type: Pseudodiphasium volubile (G. C. Order Selaginellales Prantl, Lehrb. Bot.: 116, 124. 1874.
Forst.) Holub ( Lycopodium volubile G.Forst.). Circumscription equivalent to family Selaginellaceae in this
Circumscription sensu Holub (1983). Monotypic classification. Monophyletic (Korall et al., 1999; Korall &
(Field et al., 2016). Kenrick, 2004; Zhou et al., 2015; Weststrand & Korall,
2016a). One family, one genus, and perhaps 700 species.
Pseudolycopodium Holub, Folia Geobot. Phytotax.
18: 441. 1983. Type: Pseudolycopodium densum 3. Family Selaginellaceae Willk., Anleit. Stud. Bot. 2: 163.
(Rothm.) Holub ( Lepidotis densa Rothm.). 1854. Circumscription sensu Jermy (1990b). Monophy-
Circumscription sensu Holub (1983). Monotypic letic (Korall et al., 1999; Korall & Kenrick, 2004; Zhou
(Field et al., 2016). et al., 2015; Weststrand & Korall, 2016a). One genus
and perhaps 700 species.
Spinulum A.Haines, Fam. Huperziac. Lycopodiac.
New England: 85. 2003. Type: Spinulum annotinum Selaginella P.Beauv., Mag. Encycl. 9(5): 478. 1804.
(L.) A.Haines ( Lycopodium annotinum L.). Type: Selaginella selaginoides (L.) P.Beauv. ex
Circumscription sensu Haines (2003). Presumably Schrank & Mart. ( Lycopodium selaginoides L.).
monophyletic; only one species sampled to date Circumscription sensu Jermy (1990b). Monophy-
(Field et al., 2016). Three species. letic (Korall et al., 1999; Korall & Kenrick, 2004; Zhou
et al., 2015; Weststrand & Korall, 2016a). Six (Zhou &
Subfamily Huperzioideae W.H.Wagner & Beitel ex B. Zhang, 2015) or seven (Weststrand & Korall, 2016b)
Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription subgenera can be recognized. Perhaps 700 species.
sensu Øllgaard (2015) and Field et al. (2016).
Monophyletic (Field et al., 2016). Three genera and Class Polypodiopsida Cronquist, Takht. & W.Zimm., Taxon 15:
an estimated 276 species. 133. 1966. Circumscription sensu Pryer et al. (2009) and
Ruggiero et al. (2015). Monophyletic (Pryer et al., 2001, 2004;
Huperzia Bernh., J. Bot. (Schrader) 1800(2): 126. Qiu et al., 2007; Schneider et al., 2009; Ruhfel et al., 2014;
1801. Lectotype (designated by Rothm., Repert. Wickett et al., 2014; Rothfels et al., 2015). Four subclasses,
Spec. Nov. Regni Veg. 54: 59. 1944): Huperzia selago 11 orders, 48 families, 319 genera, and an estimated 10 578
(L.) Bernh. ex Schrank & Mart. ( Lycopodium species.
selago L.). Circumscription sensu Wagner & Beitel
(1992), Zhang & Kung (1998, 2000a), Zhang & Subclass Equisetidae Warm., Osnov. Bot.: 221. 1883. Circum-
Iwatsuki (2013), Øllgaard (2015), and Field et al. scription sensu Pryer et al. (2009) and Ruggiero et al. (2015).
(2016). Monophyletic (Field et al., 2016). 25 species. Subclass consists of one extant order, one family, one genus,
and 15 species.
Phlegmariurus Holub, Preslia 36(1): 17, 21. 1964.
Type: Phlegmariurus phlegmaria (L.) T.Sen & U.Sen D. Order Equisetales DC. ex Bercht. & J.Presl, Prir. Rostlin: 271.
( Lycopodium phlegmaria L.). Circumscription 1820. Circumscription sensu Smith et al. (2006b). Order
sensu Wagner & Beitel (1992), Zhang & Kung consists of one extant family, one genus, and 15 species.
(1999, 2000b), Øllgaard (2012a, 2012b, 2015), Field &
Bostock (2013), Zhang & Iwatsuki (2013), Øllgaard & 4. Family Equisetaceae Michx. ex DC., Essai Propr. Me d.
Windisch (2014), and Field et al. (2016). Monophy- Pl.: 49. 1804. Circumscription sensu Smith et al. (2006b).
letic (Field et al., 2016). About 250 species. Family consists of one extant genus and 15 species.

PPG I 571
Equisetum L., Sp. Pl. 2: 1061. 1753. Lectotype Mankyua B.Y.Sun, M.H.Kim & C.H.Kim, Taxon 50
(designated by Hauke, Nova Hedwigia 30: 414. (4): 1020. 2002. Type: Mankyua chejuensis B.Y.Sun,
1978): Equisetum fluviatile L. Circumscription sensu M.H.Kim & C.H.Kim. Circumscription sensu Sun
Hauke (1990). Monophyletic (Guillon, 2007). An iso- et al. (2009). Monotypic (Shinohara et al., 2013).
lated genus of closely related and recently diverged
species (Des Marais et al., 2003). 15 species. Subfamily Ophioglossoideae C.Presl, Suppl. Tent.
Pterid.: 47. 1845. Circumscription sensu Hauk et al.
Subclass Ophioglossidae Klinge, Fl. Est.-Liv-Churland 1: 94. 1882. (2003). Monophyletic (Hauk et al., 2003; Shinohara
Circumscription sensu Ruggiero et al. (2015) and equivalent to et al., 2013). Four genera and an estimated 47 species.
Psilotidae in Pryer et al. (2009). Monophyletic (Pryer et al. 2001,
2004; Wickett et al., 2014; Rothfels et al., 2015). Two orders, two Cheiroglossa C.Presl, Suppl. Tent. Pterid.: 56. 1845.
families, 12 genera, and an estimated 129 species. Type: Cheiroglossa palmata (L.) C.Presl ( Ophio-
glossum palmatum L.). Circumscription sensu Hauk
E. Order Psilotales Prantl, Lehrb. Bot., ed. 5: 183. 1884. et al. (2003). Monophyletic (Hauk et al., 2003;
Circumscription sensu Smith et al. (2006b). Monophyletic Shinohara et al., 2013). Two species.
(Pryer et al., 2004; Rothfels et al., 2015). One family, two
genera, and an estimated 17 species. Ophioderma (Blume) Endl., Gen. Pl.: 66. 1836. Type:
Ophioderma pendulum (L.) C.Presl ( Ophioglos-
5. Family Psilotaceae J.W.Griff. & Henfr., Microgr. Dict.: sum pendulum L.). Circumscription sensu Hauk
540. 1855. Circumscription sensu Smith et al. (2006b). et al. (2003). Monophyletic (Hauk et al., 2003;
Monophyletic (Pryer et al., 2004). Two genera and an Shinohara et al., 2013). Three species.
estimated 17 species.
Ophioglossum L., Sp. Pl. 2: 1062. 1753. Lectotype
Psilotum Sw., J. Bot. (Schrader) 1800(2): 8, 109. (designated by J.Sm., Hist. Fil.: 369. 1875):
1801. Lectotype (designated by Brongn., Dict. Ophioglossum vulgatum L. Circumscription sensu
Class. Hist. Nat. 9: 558. 1826): Psilotum triquetrum Pichi Sermolli (1977b). Monophyletic (Hauk et al.,
Sw. Circumscription sensu Kramer (1990a). Only 2003; Shinohara et al., 2013). 41 species.
one species sampled to date, but assumed to be
monophyletic. Two species. Rhizoglossum C.Presl, Suppl. Tent. Pterid.: 47. 1845.
Type: Rhizoglossum bergianum (Schltdl.) C.Presl (
Tmesipteris Bernh., J. Bot. (Schrader) 1800(2): 8, Ophioglossum bergianum Schltdl.). Circumscription
131. 1801. Type: Tmesipteris tannensis (Spreng.) sensu Clausen (1938) and Pichi Sermolli (1977b).
Bernh. ( Lycopodium tannense Spreng.). Circum- Monotypic.
scription sensu Kramer (1990a). Monophyletic
(Perrie et al., 2010). About 15 species. Subfamily Botrychioideae C.Presl, Suppl. Tent. Pterid.:
42. 1845. Circumscription sensu Hauk et al. (2003).
F. Order Ophioglossales Link, Hort. Berol. 2: 151.1833. Monophyletic (Hauk et al., 2003; Shinohara et al.,
Circumscription sensu Smith et al. (2006b). Monophyletic 2013). Four genera and an estimated 63 species.
(Pryer et al., 2004; Rothfels et al., 2015). One family, ten
genera, and an estimated 112 species. Botrychium Sw., J. Bot. (Schrader) 1800(2): 110.
1801. Lectotype (designated by J.Sm., Hist. Fil.:
6. Family Ophioglossaceae Martinov, Tekhno-Bot. Slo- 369. 1875): Botrychium lunaria (L.) Sw. ( Osmunda
var: 438. 1820. Circumscription sensu Smith et al. lunaria L.). Circumscription sensu Wagner (1990)
(2006b). Monophyletic (Pryer et al., 2004; Rothfels and Dauphin et al. (2014). Monophyletic (Hauk
et al., 2015). Four subfamilies, ten genera, and an et al., 2003). About 35 species.
Botrypus Michx., Fl. Bor.-Amer. (Michaux) 2: 274.
Subfamily Helminthostachyoideae C.Presl, Suppl. Tent. 1803. Lectotype (designated by Pic.Serm., Webbia
Pterid.: 58. 1845. Circumscription sensu Shinohara et al. 26: 497. 1972): Botrypus virginianus (L.) Michx. (
(2013). Subfamily consists of a single monotypic genus. Osmunda virginiana L.). Circumscription sensu
Hauk et al. (2003). Includes: Osmundopteris (Milde)
Helminthostachys Kaulf., Enum. Filic.: 28. 1824. Small. Not monophyletic (Hauk et al., 2003;
Type: Helminthostachys zeylanica (L.) Hook. ( Shinohara et al., 2013). About two species.
Osmunda zeylanica L.). Circumscription sensu Wag-
ner (1990). Includes: Ophiala Desv. Monotypic Japanobotrychium Masam., Journ. Soc. Trop.
(Hauk et al., 2003; Shinohara et al., 2013). Agric. Formosa 3: 246. 1931. Type: Japanobo-
trychium arisanense Masam. Monotypic (Shinohara
Subfamily Mankyuoideae J.R.Grant & B.Dauphin, this et al., 2013).
classification (see below). Circumscription sensu
Shinohara et al. (2013). Subfamily consists of a single Sceptridium Lyon, Bot. Gaz. 40(6): 457. 1905.
monotypic genus. Lectotype (designated by R.T.Clausen, Mem.

Torrey Bot. Club 19(2): 24. 1938): Sceptridium Subclass Polypodiidae Cronquist, Takht. & W.Zimm., Taxon 15:
obliquum (Muhl. ex Willd.) Lyon ( Botrychium 133. 1966. Circumscription sensu Pryer et al. (2008) and
obliquum Muhl. ex Willd.). Circumscription sensu Ruggiero et al. (2015). Monophyletic (Pryer et al., 2004;
Hauk et al. (2003). Monophyletic (Hauk et al., Rothfels et al., 2015). Seven orders, 44 families, 300 genera,
2003). About 25 species. and an estimated 10 323 species.
Subclass Marattiidae Klinge, Fl. Est.-Liv-Churland 1: 93. 1882. H. Order Osmundales Link, Hort. Berol.: 445. 1833. Circum-
Circumscription sensu Christenhusz (2009) and Ruggiero et al. scription sensu Smith et al. (2006b). Order consists of one
(2015). Monophyletic (Pryer et al., 2004). One order, one family. One family, six genera, and an estimated 18 species.
family, six genera, and an estimated 111 species.
8. Family Osmundaceae Martinov, Tekhno-Bot. Slovar.:
G. Order Marattiales Link, Hort. Berol. 2: 148. 1833. 445. 1820. Circumscription sensu Smith et al. (2006b).
Circumscription sensu Smith et al. (2006b). Monophyletic Monophyletic (Yatabe et al., 1999; Schuettpelz &
(Pryer et al., 2004). One family, six genera, and an Pryer, 2007; Metzgar et al., 2008). Six genera and an
estimated 111 species. estimated 18 species.
7. Family Marattiaceae Kaulf., Enum. Filic.: 31. 1824. Cir- Claytosmunda (Y.Yatabe, N.Murak. & K.Iwats.)
cumscription sensu Smith et al. (2006b). Monophyletic Metzgar & Rouhan, this classification (see below).
(Pryer et al., 2004). Six genera and an estimated 111 Type: Claytosmunda claytoniana (L.) Metzgar &
species. Rouhan ( Osmunda claytoniana L.). Circumscrip-
tion equivalent to Osmunda subgenus Claytos-
Angiopteris Hoffm., Commentat. Soc. Regiae Sci. munda in Yatabe et al. (2005). Monotypic.
Gott. 12: 29. 1796. Type: Angiopteris evecta
(G.Forst.) Hoffm. ( Polypodium evectum G.Forst.). Leptopteris C.Presl, Suppl. Tent. Pterid.: 70. 1845.
Circumscription sensu Murdock (2008). Includes: Lectotype (designated by C.Chr., Index Filic.
Archangiopteris Christ & Giesenh.; Clementea lvi.1906): Leptopteris hymenophylloides (A.Rich.)
Cav.; Macroglossum Copel.; Protangiopteris C.Presl ( Todea hymenophylloides A.Rich.). Cir-
Hayata; Protomarattia Hayata; Psilodochea C.Presl. cumscription sensu Kramer (1990o). Monophyletic
Monophyletic (Murdock, 2008). Perhaps 30 (Yatabe et al., 1999; Metzgar et al., 2008; Bomfleur
species. et al., 2015). Six species.
Christensenia Maxon, Proc. Biol. Soc. Washington 18 Osmunda L., Sp. Pl. 2: 1063. 1753. Lectotype
(50): 239. 1905. Type: Christensenia aesculifolia (designated by Leman, Dict. Sci. Nat. 37: 9. 1825):
(Blume) Maxon ( Aspidium aesculifolium Blume). Osmunda regalis L. Circumscription equivalent to
Circumscription sensu Murdock (2008). Includes: subgenus Osmunda in Kramer (1990o). Monophy-
Kaulfussia Blume.; Macrostoma Griff. Only one letic (Yatabe et al., 1999; Metzgar et al., 2008;
species sampled to date, but assumed to be Bomfleur et al., 2015). About four species.
monophyletic (Murdock, 2008). Two species.
Osmundastrum C.Presl, Gef€assb€ undel Farrn: 18.
Danaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 420, 1847. Type: Osmundastrum cinnamomeum (L.) C.
pl.9, f.11. 1793. Type: Danaea nodosa (L.) Sm. ( Presl ( Osmunda cinnamomea L.). Circumscription
Acrostichum nodosum L.). Circumscription sensu sensu Yatabe et al. (2005). Monotypic (Miller, 1967;
Murdock (2008). Includes: Heterodanaea C.Presl. Yatabe et al., 1999; Metzgar et al., 2008).
Monophyletic (Christenhusz et al., 2008; Murdock,
2008). Perhaps 50 species. Plenasium C.Presl, Tent. Pterid.: 109, pl.3, f.13. 1836.
Lectotype (designated by C.Chr., Index Filic. lvi.
Eupodium J.Sm. in Hooker, Gen. Fil. (Hooker): t.118. 1906): Plenasium banksiaefolium (C.Presl) C.Presl.
1842. Type: Eupodium kaulfussii (J.Sm.) J.Sm. ( ( Nephrodium banksiaefolium C.Presl). Circum-
Marattia kaulfussii J.Sm.). Circumscription sensu scription equivalent to subgenus Plenasium in
Murdock (2008). Monophyletic (Murdock, 2008). Kramer (1990o). Monophyletic (Yatabe et al.,
Two species. 1999; Metzgar et al., 2008; Bomfleur et al., 2015).
About four species.
Marattia Sw., Prodr. 8: 128. 1788. Type: Marattia
alata Sw. Circumscription sensu Murdock (2008). Todea Willd. ex Bernh., J. Bot. (Schrader) 1800(2):
Includes: Gymnotheca C.Presl; Stibasia C.Presl. 126. 1801. Type: Todea africana Willd. ex Bernh.,
Monophyletic (Murdock, 2008). Seven species. nom illeg. superfl. ( Acrostichum barbarum L.).
Circumscription sensu Kramer (1990o). Monophy-
Ptisana Murdock, Taxon 57(3): 744. 2008. Type: letic (Metzgar et al., 2008). Two species.
Ptisana salicina (Sm.) Murdock ( Marattia salicina
Sm.). Circumscription sensu Murdock (2008). I. Order Hymenophyllales A.B.Frank, Syn. Pflanzenk., ed. 2,
Monophyletic (Murdock, 2008). 20 species. 3: 1452. 1877. Circumscription sensu Smith et al. (2006b).

PPG I 573
Order consists of one family, nine genera, and an Trichomanes radicans Sw.). Circumscription sensu
estimated 434 species. Ebihara et al. (2006). Monophyletic (Ebihara et al.,
2007). About 15 species.
9. Family Hymenophyllaceae Mart., Consp. Regn. Veg.: 3.
1835. Circumscription sensu Smith et al. (2006b). Subfamily Hymenophylloideae Burnett, Outlines Bot.:
Monophyletic (Schuettpelz & Pryer, 2007). Two 324. 1835. Circumscription equivalent to “hymenophyl-
subfamilies, nine genera, and an estimated 434 loids” in Ebihara et al. (2006). Subfamily consists of one
species. genus and an estimated 250 species.
Subfamily Trichomanoideae C.Presl, Hymenophylla- Hymenophyllum Sm., Me m. Acad. Roy. Sci. (Turin)
ceae: 102. 1843. Circumscription equivalent to “tricho- 5: 418, pl.9, f.8. 1793. Lectotype (designated by
manoids” in Ebihara et al. (2006). Monophyletic (Pryer C.Presl, Hymenophyllaceae: 1. 1844): Hymenophyl-
et al., 2001; Dubuisson et al., 2003; Ebihara et al., 2007). lum tunbrigense (L.) Sm. ( Trichomanes tunbri-
Relationships among the trichomanoid genera are still gense L.). Circumscription sensu Ebihara et al.
mostly unresolved. Eight genera and an estimated 184 (2006). Monophyletic (Pryer et al., 2001; Hennequin
species. et al., 2006; Ebihara et al., 2007). About 250
species.
Abrodictyum C.Presl, Hymenophyllaceae: 20. 1843.
Type: Abrodictyum cumingii C.Presl. Circumscrip- J. Order Gleicheniales Schimp., Traite Paleont. Veg. 1: 669.
tion sensu Ebihara et al. (2006). Monophyletic 1869. Circumscription sensu Smith et al. (2006b). Mono-
(Ebihara et al., 2007). About 25 species. phyletic (Pryer et al., 2004). Three families, ten genera, and
an estimated 172 species.
Callistopteris Copel., Philipp. J. Sci. 67: 49. 1938.
Type: Callistopteris apiifolia (C.Presl) Copel. ( 10. Family Matoniaceae C.Presl, Gef€assb€
undel Farrn: 32.
Trichomanes apiifolium C.Presl). Circumscription 1847. Circumscription sensu Smith et al. (2006b).
sensu Ebihara et al. (2006). Monophyletic (Ebihara Monophyletic (Kato & Setoguchi, 1998). Two genera
et al., 2007). Five species. and an estimated four species.
Cephalomanes C.Presl, Hymenophyllaceae: 17. Matonia R.Br. ex Wall., Pl. Asiat. Rar. 1(1): 16. 1829.
1843. Type: Cephalomanes atrovirens C.Presl. Type: Matonia pectinata R.Br. Circumscription
Circumscription sensu Ebihara et al. (2006). sensu Kramer (1990l). Monophyletic (Kato &
Monophyletic (Ebihara et al., 2007). Four species. Setoguchi, 1998). Two species.
Crepidomanes C.Presl, Epimel. Bot.: 258. 1851. Phanerosorus Copel., Philipp. J. Sci. C 3(6): 344.
Type: Crepidomanes intramarginale (Hook. & 1909. Type: Phanerosorus sarmentosus (Baker)
Grev.) C.Presl ( Trichomanes intramarginale Copel. ( Matonia sarmentosa Baker). Circumscrip-
Hook. & Grev.). Circumscription sensu Ebihara tion sensu Kramer (1990l). Monophyletic (Kato &
et al. (2006). Includes: Nesopteris Copel. Mono- Setoguchi, 1998). Two species.
phyletic (Ebihara et al., 2007). About 30 species.
11. Family Dipteridaceae Seward & E.Dale, Philos. Trans.,
Didymoglossum Desv., M em. Soc. Linn. Paris 6: 330. Ser. B. 194: 487, 499, 502. 1901. Circumscription
1827. Lectotype (designated by C.Chr., Index Filic. sensu Smith et al. (2006b). Monophyletic (Pryer
xiv. 1906): Didymoglossum hymenoides (Hedw.) et al., 2004). Two genera and an estimated 11
Copel. ( Trichomanes hymenoides Hedw.). Circum- species.
scription sensu Ebihara et al. (2006). Monophyletic
(Ebihara et al., 2007). About 30 species. Cheiropleuria C.Presl, Epimel. Bot.: 189. 1851.
Lectotype (designated by J.Sm., Hist. Fil.: 139.
Polyphlebium Copel., Philipp. J. Sci. 67: 55. 1938. 1875): Cheiropleuria bicuspis (Blume) C.Presl (
Type: Polyphlebium venosum (R.Br.) Copel. ( Polypodium bicuspe Blume). Circumscription
Trichomanes venosum R.Br.). Circumscription sensu sensu Kramer (1990b). Monophyletic (Kato
Ebihara et al. (2006). Monophyletic (Ebihara et al., et al., 2001). Three species.
Dipteris Reinw., Syll. Pl. Nov. 2: 3. 1825. Type:
Trichomanes L., Sp. Pl. 2: 1097. 1753. Lectotype Dipteris conjugata Reinw. Circumscription sensu
(designated by Underw., Mem. Torrey Bot. Club 6: Kramer (1990f). Presumably monophyletic (sam-
256, 283. 1899): Trichomanes crispum L. Circum- pling limited to date). About eight species.
scription sensu Ebihara et al. (2006). Monophyletic
(Ebihara et al., 2007). About 60 species. 12. Family Gleicheniaceae C.Presl, Reliq. Haenk. 1: 70.
1825. Circumscription sensu Smith et al. (2006b).
Vandenboschia Copel., Philipp. J. Sci. 67: 51. 1938. Monophyletic (Pryer et al., 2004). Six genera and an
Type: Vandenboschia radicans (Sw.) Copel. ( estimated 157 species.

Dicranopteris Bernh., Neues J. Bot. 1(2): 38–39. Monophyletic (Wikstr€

om et al., 2002). Two genera and
1805. Type: Dicranopteris dichotoma (Thunb.) an estimated 35 species.
Bernh. ( Polypodium dichotomum Thunb.).
Circumscription sensu Kramer (1990g), but ex- Actinostachys Wall., Numer. List 1. 1829. Type:
cluding Gleichenella. Sampling to date has been Actinostachys digitata (L.) Wall. ( Acrostichum
insufficient to fully assess the monophyly of the digitatum L.). Circumscription sensu Mickel &
genus (Li et al., 2010). Perhaps 20 species. Smith (2004). Monophyletic (Wikstr€ om et al.,
Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus.
Tokyo 29: 47. 1950. Lectotype (designated by C. Schizaea Sm., Me m. Acad. Roy. Sci. (Turin) 5: 419,
Chr., Index Filic. liv. 1906): Diplopterygium pl.9, f.9. 1793. Type: Schizaea dichotoma (L.) J.Sm.
glaucum (Thunb. ex Houtt.) Nakai ( Polypodium ( Acrostichum dichotomum L.). Circumscription
glaucum Thunb. ex Houtt.). Circumscription sensu Mickel & Smith (2004). Monophyletic
sensu Kramer (1990g). Sampling to date has (Wikstro €m et al., 2002). Perhaps 20 species.
been insufficient to fully assess the monophyly of
the genus (Li et al., 2010). About 25 species. 15. Family Anemiaceae Link, Fil. Spec.: 23. 1841. Circum-
scription sensu Smith et al. (2006b). Monophyletic
Gleichenella Ching, Sunyatsenia 5: 276. 1940. (Labiak et al., 2015a). One genus and an estimated 115
Type: Gleichenella pectinata (Willd.) Ching ( species.
Mertensia pectinata Willd.). Circumscription sensu
Mickel & Smith (2004). Monotypic. Anemia Sw., Syn. Fil.: 6, 155. 1806. Lectotype
(designated by Underw., Mem. Torrey Bot. Club 6:
Gleichenia Sm., Mem. Acad. Roy. Sci. (Turin) 5: 267, 276. 1899): Anemia phyllitidis (L.) Sw. (
419, pl.9, f.10. 1793. Type: Gleichenia polypo- Osmunda phyllitidis L.). Circumscription sensu
dioides (L.) Sm. ( Onoclea polypodioides L.). Mickel (2016). Includes: Mohria Sw.; Colina Greene.
Circumscription sensu Kramer (1990g). Appar- Monophyletic (Labiak et al., 2015a). About 115
ently monophyletic (Perrie et al., 2007; but see species.
Hennequin et al., 2014), although sampling to
date has been limited. About 15 species. L. Order Salviniales Link, Hort. Berol. 3: 155. 1833. Circum-
scription sensu Smith et al. (2006b). Monophyletic (Pryer
Sticherus C.Presl, Tent. Pterid.: 51. 1836. Lecto- et al., 1995; Pryer, 1999; Schuettpelz & Pryer, 2007;
type (designated by C.Chr., Index Filic. liv. 1906): Rothfels et al., 2015). Two families, five genera, and an
Sticherus laevigatus (Willd.) C.Presl ( Mertensia estimated 82 species.
laevigata Willd.). Circumscription sensu Kramer
(1990g). Apparently monophyletic (Perrie et al., 16. Family Salviniaceae Martinov, Tekhno-Bot.Slovar.:
2007), although sampling to date has been 559. 1820. Circumscription sensu Smith et al.
limited. About 95 species. (2006b). Monophyletic (Pryer et al., 1995; Pryer,
1999; Schuettpelz & Pryer, 2007; Nagalingum et al.,
Stromatopteris Mett., Ann. Sci. Nat. Bot., ser. 4, 2008). Two genera and an estimated 21 species.
15: 84. 1861. Type: Stromatopteris moniliformis
Mett. Circumscription sensu Kramer (1990g). Azolla Lam., Encycl. 1(1): 343. 1783. Type: Azolla
Monotypic. filiculoides Lam. Circumscription sensu Schneller
(1990a). Monophyletic (Metzgar et al., 2007; Reid
K. Order Schizaeales Schimp., Traite Paleont. Ve
g. 1: 674. et al., 2006; Nagalingum et al., 2008). About nine
1869. Circumscription sensu Smith et al. (2006b). Mono- species.
phyletic (Schuettpelz & Pryer, 2007). Three families, four
genera, and an estimated 190 species. g., Pl. Veron. 3: 52. 1754. Type: Salvinia
Salvinia Se
natans (L.) All. ( Marsilea natans L.). Circumscrip-
13. Family Lygodiaceae M.Roem., Handb. Allg. Bot. 3: 520. tion sensu Schneller (1990b). Monophyletic (Na-
1840. Circumscription sensu Smith et al. (2006b). galingum et al., 2008). About 12 species.
Monophyletic (Wikstro€ m et al., 2002). One genus and
an estimated 40 species. 17. Family Marsileaceae Mirb., Hist. Nat. Veg. 5: 126. 1802.
Circumscription sensu Smith et al. (2006b). Monophy-
Lygodium Sw., J. Bot. (Schrader) 1800(2): 7, 106. letic (Pryer, 1999; Schuettpelz & Pryer, 2007; Naga-
1801. Type: Lygodium scandens (L.) Sw. ( lingum et al., 2008). Three genera and an estimated 61
Ophioglossum scandens L.). Circumscription sensu species.
Kramer (1990q). Monophyletic (Wikstro €m et al.,
2002). Perhaps 40 species. Marsilea L., Sp. Pl. 2: 1099. 1753. Lectotype
(designated by Maxon, Sci. Surv. Porto Rico &
14. Family Schizaeaceae Kaulf., Wesen Farrenkr.: 119. Virgin Islands 6(3): 509. 1926): Marsilea quadrifolia
1827. Circumscription sensu Smith et al. (2006b). L. Circumscription sensu Kramer (1990k).

PPG I 575
Monophyletic (Nagalingum et al., 2007, 2008; Presumably monophyletic; seven species sampled
Whitten et al., 2012). About 55 species. to date (Korall et al., 2006; Ebihara, 2011). 15
species.
Pilularia L., Sp. Pl. 2: 1100. 1753. Type: Pilularia
globulifera L. Circumscription sensu Kramer 22. Family Cibotiaceae Korall, Taxon 55(3): 712. 2006. Cir-
(1990k). Monophyletic (Nagalingum et al., 2008). cumscription sensu Smith et al. (2006b). Family con-
About five species. sists of a single genus and an estimated nine species.
Regnellidium Lindm., Ark. Bot. 3(6): 2. 1904. Type: Cibotium Kaulf., Berlin. Jahrb. Pharm. Verbunde-
Regnellidium diphyllum Lindm. Circumscription nen Wiss. 21: 53. 1820. Type: Cibotium chamissoi
sensu Kramer (1990k). Monotypic. Kaulf. Circumscription sensu Kramer (1990e).
Includes: Pinonia Gaudich. Monophyletic (Geiger
M. Order Cyatheales A.B.Frank in Leunis, Syn. Pflanzenk. (ed. et al., 2013). Nine species.
2) 3: 1452. 1877. Circumscription sensu Smith et al. (2006b).
Monophyletic (Korall et al., 2006; Schuettpelz et al., 2007; 23. Family Metaxyaceae Pic.Serm., Webbia 24(2): 701.
Rothfels et al., 2015). Eight families, 13 genera, and an 1970. Circumscription sensu Smith et al. (2006b).
estimated 713 species. Family consists of a single genus and an estimated six
species.
18. Family Thyrsopteridaceae C.Presl, Gef€assb€
undel Farrn:
22, 38. 1847. Circumscription sensu Smith et al. Metaxya C.Presl, Tent. Pterid.: 59. 1836. Type:
(2006b). Family consists of a single monotypic genus. Metaxya rostrata (Kunth) C.Presl ( Polypodium
rostratum Humb. & Bonpl. ex Willd.). Circumscrip-
Thyrsopteris Kunze, Linnaea 9: 507. 1835. Type: tion sensu Cardenas et al. (2016). Includes:
Thyrsopteris elegans Kunze. Circumscription sensu Amphidesmium J.Sm.; Amphidesmium Schott ex
Kramer (1990e). Includes: Panicularia Colla. Mono- Kunze. Monophyletic (Cardenas et al., 2016). Six
typic (Korall et al., 2006). species.
19. Family Loxsomataceae C.Presl, Gef€assb€

undel Farrn: 31. 24. Family Dicksoniaceae M.R.Schomb., Reis. Br.-Guiana 2:
1847. Circumscription sensu Smith et al. (2006b). 1047. 1849. Circumscription sensu Smith et al. (2006b).
Monophyletic (Korall et al., 2006). Two monotypic Monophyletic (Korall et al., 2006; Adjie & Lestari, 2014;
genera. Noben et al., pers. comm.). Three genera and an
Loxsoma R.Br. ex Hook. in A.Cunn., Companion
Bot. Mag. 2: 366. 1837. Type: Loxsoma cunninghamii Calochlaena (Maxon) M.D.Turner & R.A.White,
R.Br. ex Hook. Circumscription sensu Kramer Amer. Fern J. 78(3): 91. 1988. Type: Culcita dubia
(1990j). Includes: Chrysodium F ee. Monotypic. (R.Br.) Maxon ( Davallia dubia R.Br.). Circum-
scription sensu White & Turner (1988). Monophy-
Loxsomopsis Christ, Bull. Herb. Boissier, ser. 2, 4(5): letic (Adjie & Lestari, 2014; Noben et al., pers.
399. 1904. Type: Loxsomopsis costaricensis Christ. comm.). Five species.
Circumscription sensu Lehnert et al. (2001).
Monotypic. Dicksonia L’Her., Sert. Angl. 30. 1789. Lectotype
(designated by J.Sm., Hist. Fil.: 254. 1875): Dicksonia
20. Family Culcitaceae Pic.Serm., Webbia 24: 702. 1970. arborescens L’Her. Circumscription sensu Kramer
Circumscription sensu Smith et al. (2006b). Monophy- (1990e). Includes: Balantium Kaulf. Monophyletic
letic (Korall et al., 2006). One genus and two species. (Noben et al., pers. comm.). 26 species.
Culcita C.Presl, Tent. Pterid.: 135, t.5, f.5. 1836. Lophosoria C.Presl, Gef€assb€
undel Farrn: 36. 1847.
Type: Culcita macrocarpa C.Presl. Circumscription Lectotype (designated by J.Sm., Hist. Fil.: 251.
sensu White & Turner (1988). Monophyletic (Korall 1875): Lophosoria pruinata (Sw.) C.Presl ( Poly-
et al., 2006). Two species. podium pruinatum Sw.). Circumscription sensu
Kramer (1990i). Includes: Trichosorus Liebm.
21. Family Plagiogyriaceae Bower, Ann. Bot. (Oxford) 40: Monophyletic (Noben et al., pers. comm.). About
484. 1926. Circumscription sensu Smith et al. (2006b). four species.
Family consists of a single genus and an estimated 15
species. 25. Family Cyatheaceae Kaulf., Wesen Farrenkr.: 119. 1827.
Circumscription sensu Smith et al. (2006b). Monophy-
Plagiogyria (Kunze) Mett., Abh. Senckenberg. letic (Korall et al., 2006, 2007). Three genera and an
Naturf. Ges. 2: 1, 268. 1858. Lectotype (designated estimated 643 species.
by C.Chr., Index Filic. xliii. 1906): Plagiogyria
euphlebia (Kunze) Mett. ( Lomaria euphlebia Alsophila R.Br., Prodr.: 158. 1810. Type: Alsophila
Kunze). Circumscription sensu Kramer (1990p). australis R.Br. Circumscription sensu Conant

(1983). Includes: Amphicosmia Gardner; Dichorexia sorbifolia Sm.). Circumscription sensu Kramer
C.Presl; Gymnosphaera Blume; Nephelea R.M. (1990e). Monotypic.
Tryon; Thysanobotrya Alderw. Monophyletic (Kor-
all et al., 2007). 275 species. 28. Family Lonchitidaceae Doweld, New Syllab. Pl. Fam.:
353. 2006. Circumscription sensu Christenhusz et al.
Cyathea Sm., M em. Acad. Roy. Sci. (Turin) 5: 416. (2011). Family consists of a single genus and an
1793. Lectotype (designated by J.Sm., Hist. Fil.: estimated two species.
244. 1875): Cyathea arborea (L.) Sm. ( Polypodium
arboreum L.). Circumscription sensu Korall et al. Lonchitis L., Sp. Pl. 2: 1078. 1753. Lectotype
(2007). Includes: Actinophlebia C.Presl; Chnoo- (designated by Brongn., Dict. Class. Hist. Nat. 9:
phora Kaulf.; Cnemidaria C.Presl; Cnemidopteris 490. 1826): Lonchitis hirsuta L. Circumscription
Reichenb.; Cormophyllum Newm.; Disphenia C. sensu Kramer (1990d). Presumably monophyletic,
Presl; Hemitelia R.Br.; Hemistegia C.Presl; Hyme- but only one species sampled to date. An
nophyllopsis K.I.Goebel; Microstegnus C.Presl; estimated two species.
Trichopteris C.Presl; Cyathidaria Caluff. Mono-
phyletic (Korall et al., 2007). 265 species. 29. Family Lindsaeaceae C.Presl ex M.R.Schomb., Reis.
Br.-Guiana 3: 883. 1849. Circumscription sensu Chris-
Sphaeropteris Bernh., J. Bot. (Schrader) 1800(2): tenhusz et al. (2011). Monophyletic (Lehtonen
122. 1801. Type: Sphaeropteris medullaris (G.Forst.) et al., 2010). Seven genera and an estimated 234
Bernh. ( Polypodium medullare G.Forst.). Circum- species.
scription sensu Tryon (1970). Includes: Eatoniop-
teris Bomm.; Fourniera Bomm.; Schizocaena J.Sm. Lindsaea Dryand. ex Sm., Me m. Acad. Roy. Sci.
Monophyletic (Korall et al., 2007). 103 species. (Turin) 5: 413, pl.9, f.4. 1793. Lectotype (designated
by Maxon, Sci. Surv. Porto Rico & Virgin Islands 6
N. Order Polypodiales Link, Hort. Berol. 2: 5. 1833. Circum- (3): 488. 1926): Lindsaea guianensis (Aubl.) Dryand.
scription sensu Smith et al. (2006b). Monophyletic ( Adiantum guianense Aubl.). Circumscription
(Schuettpelz & Pryer, 2007). Six suborders, 26 families, sensu Lehtonen et al. (2010). Includes: Guerinia J.
253 genera, and an estimated 8714 species. Sm.; Humblotiella Tardieu; Lindsaenium Fe e; Odon-
toloma Maxon; Sambirania Tardieu; Schizolegnia
Suborder Saccolomatineae Hovenkamp, this classification Alston; Schizoloma Gaudich.; Synaphlebium J.Sm.
(see below). Circumscription equivalent to family Sacco- Monophyletic (Lehtonen et al., 2010). About 180
lomataceae in this classification and in Smith et al. (2006b). species.
Additional investigation is needed to establish monophyly.
One family, one genus, and an estimated 18 species. Nesolindsaea Lehtonen & Christenh., Bot. J. Linn.
Soc. 163(3): 336–337. 2010. Type: Nesolindsaea
26. Family Saccolomataceae Doweld, Phytologia 90(3): caudata (Hook.) Lehtonen & Christenh. ( Lind-
417. 2008. Circumscription sensu Smith et al. (2006b). saea caudata Hook.). Circumscription sensu Lehto-
Contains a single genus, but additional investigation is nen et al. (2010). Monophyletic (Lehtonen et al.,
needed (see below). An estimated 18 species. 2010). Two species.
Saccoloma Kaulf., Berlin. Jahrb. Pharm. Verbunde- Odontosoria Fee, M em. Foug., 5. Gen. Filic. 325.
nen Wiss. 21: 51. 1820. Type: Saccoloma elegans 1852. Type: Odontosoria uncinella (Kunze) F ee (
Kaulf. Circumscription sensu Kramer (1990d). Inclu- Davallia uncinella Kunze). Circumscription sensu
des: Orthiopteris Copel. Additional investigation is Lehtonen et al. (2010). Includes: Lindsayopsis Kuhn.
needed (Perrie et al., 2015), as indications that the Monophyletic (Lehtonen et al., 2010). About 23
genus is not monophyletic may be based on errors species.
in sequencing or mixed samples. 18 species.
Osmolindsaea (K.U.Kramer) Lehtonen & Chris-
Suborder Lindsaeineae Lehtonen & Tuomisto, this classifi- tenh., Bot. J. Linn. Soc. 163(3): 335. 2010. Type:
cation (see below). Circumscription equivalent to Lind- Osmolindsaea odorata (Roxb.) Lehtonen & Chris-
saeaceae in Smith et al. (2006b). Probably monophyletic, tenh. ( Lindsaea odorata Roxb.). Circumscrip-
but needs to be confirmed with additional data (Lehtonen tion sensu Lehtonen et al. (2010, 2013).
et al., 2012). Three families, nine genera, and an estimated Monophyletic (Lehtonen et al., 2010, 2013). About
237 species. seven species.
27. Family Cystodiaceae J.R.Croft, Kew Bull. 41(4): 797. Sphenomeris Maxon, J. Wash. Acad. Sci. 3(5): 144.
1986. Circumscription sensu Christenhusz et al. (2011). 1913. Type: Sphenomeris clavata (L.) Maxon (
Family consists of a single monotypic genus. Adiantum clavatum L.). Circumscription sensu
Lehtonen et al. (2010). Only one species sampled
Cystodium J.Sm, Gen. Fil. (Hooker) t.96. 1841. Type: to date, but assumed to be monophyletic. Three
Cystodium sorbifolium (Sm.) J.Sm. ( Dicksonia species.

PPG I 577
Tapeinidium (C.Presl) C.Chr., Index Filic.: 631. 1906. Jessen et al. (2012). Monophyletic (Metzgar et al.,
Lectotype (designated by Copel., Ann. Cryptog. 2013). About 10 species.
Phytopathol. 5 [Gen. Fil.]: 53. 1947): Tapeinidium
pinnatum (Cav.) C.Chr. ( Davallia pinnata Cav.). Llavea Lag., Gen. Sp. Pl.: 33. 1816. Type: Llavea
Circumscription sensu Lehtonen et al. (2010). cordifolia Lag. Circumscription sensu Tryon et al.
Includes: Protolindsaya Copel. Monophyletic (Leh- (1990). Monotypic.
tonen et al., 2010). 18 species.
Subfamily Pteridoideae Link, Fil. Spec.: 48. 1841.
Xyropteris K.U.Kramer, Acta Bot. Neerl. 6: 599. Circumscription sensu Zhang et al. (2015). Monophy-
1958. Type: Xyropteris stortii (Alderw.) K.U.Kramer letic (Schuettpelz et al., 2007; Zhang et al., 2015). 13
( Schizoloma stortii Alderw.). Circumscription genera and an estimated 400 species.
sensu Lehtonen et al. (2010). Monotypic.
Actiniopteris Link, Fil. Spec.: 79. 1841. Lectotype
Suborder Pteridineae J.Prado & Schuettp., this classifica- (designated by Pic.Serm., Webbia 17: 6–7. 1962):
tion (see below). Circumscription equivalent to family Actiniopteris radiata (Sw.) Link ( Asplenium
Pteridaceae in this classification and in Smith et al. radiatum Sw.). Circumscription sensu Tryon et al.
(2006b). Monophyletic (Schuettpelz & Pryer, 2007; (1990). Monophyletic (Schneider et al., 2013). Six
Rothfels et al., 2015). One family, 53 genera, and an species.
Anogramma Link, Fil. Spec.: 137. 1841. Lectotype
30. Family Pteridaceae E.D.M.Kirchn., Schul-Bot.: 109. (designated by C.Chr., Index Filic. xxxvii. 1906):
1831. Circumscription sensu Smith et al. (2006b). Anogramma leptophylla (L.) Link ( Polypodium
Monophyletic (Schuettpelz & Pryer, 2007). Five leptophyllum L.). Circumscription sensu Nakazato
subfamilies, 53 genera, and an estimated 1211 species. & Gastony (2003). Polyphyletic (Schneider et al.,
2013; Hennequin et al., 2014). About five species.
Subfamily Parkerioideae Burnett, Outlines Bot.: 324.
1835, as “Parkeridae”. Circumscription equivalent to Austrogramme E.Fourn., Ann. Sci. Nat. Bot., ser. 5,
“ceratopteridoids” in Schuettpelz et al. (2007). 18: 278. 1873. Lectotype (designated by C.Chr.,
Monophyletic (Schuettpelz et al., 2007). Two genera Index Filic. xxxvii. 1906): Austrogramme marginata
and an estimated nine species. (Mett.) E.Fourn. ( Gymnogramma marginata
Mett.). Circumscription sensu Tryon et al. (1990).
Acrostichum L., Sp. Pl. 2: 1067. 1753. Lectotype Monophyletic (Cochran et al., 2014), but relatively
(designated by J.Sm., Hist. Fil.: 146. 1875): Acros- few species sampled. Six species.
tichum aureum L. Circumscription sensu Tryon
et al. (1990). Monophyletic (Schuettpelz et al., Cerosora Domin, Acta Bot. Bohem. 8: 3. 1929. Type:
2007). Three species. Cerosora chrysosora (Baker) Domin ( Gymnog-
ramma chrysosora Baker). Circumscription sensu
Ceratopteris Brongn., Bull. Sci. Soc. Philom. Paris Schneider et al. (2013). Monophyletic (Schneider
1821: 186. 1822. Lectotype (designated by Brongn., et al., 2013). Four species.
Dict. Class. Hist. Nat. 3: 350. 1823): Ceratopteris
thalictroides (L.) Brongn. ( Acrostichum thalic- Cosentinia Tod., Giorn. Sci. Nat. Econ. Palermo 1:
troides L.). Circumscription sensu Tryon et al. 219. 1866. Type: Cosentinia vellea (Aiton) Tod. (
(1990). Monophyletic (Schuettpelz et al., 2007). Acrostichum velleum Aiton). Circumscription sensu
Perhaps six species. Tryon et al. (1990). Monotypic.
Subfamily Cryptogrammoideae S.Lindsay, Edinburgh. Jamesonia Hook. & Grev., Pl. Voy. Russes Monde 2:
J. Bot. 66(2): 358. 2009. Circumscription equivalent to t.178. 1830. Type: Jamesonia pulchra Hook. & Grev.
“cryptogrammoids” in Schuettpelz et al. (2007). Circumscription sensu Pabo n-Mora & Gonzalez
Monophyletic (Schuettpelz et al., 2007; Metzgar e; Nephopteris Lellin-
(2015). Includes: Eriosorus Fe
et al., 2013). Three genera and an estimated 31 ger. Monophyletic (Pabo n-Mora & Gonzalez, 2015).
species. About 50 species.
Coniogramme Fe e, Mem. Foug., 5. Gen. Filic.: 167. Onychium Kaulf., Berlin. Jahrb. Pharm. Verbunde-
1852. Type: Coniogramme javanica (Blume) Fe e ( nen Wiss. 21: 45. 1820. Type: Onychium capense
Gymnogramma javanica Blume). Circumscription Kaulf. Circumscription sensu Tryon et al. (1990).
sensu Tryon et al. (1990). Monophyletic (Metzgar Monophyletic (Schuettpelz et al., 2007). Perhaps
et al., 2013). At least 20 species. 10 species.
Cryptogramma R.Br., Narr. Journey Polar Sea: 767. Pityrogramma Link, Handbuch 3: 19–20. 1833.
1823. Type: Cryptogramma acrostichoides R.Br. Type: Pityrogramma chrysophylla (Sw.) Link (
Circumscription sensu Metzgar et al. (2013) and Acrostichum chrysophyllum Sw.). Circumscription

sensu Tryon et al. (1990). Includes: Trismeria F

ee. Schuettp. ( Hemionitis boryana Willd.). Circum-
Presumably monophyletic, but sampling incom- scription sensu Schuettpelz et al. (2016). Mono-
plete (Schneider et al., 2013; Zhang et al., 2015). phyletic (Schuettpelz et al., 2016). Three species.
About 20 species.
Antrophyum Kaulf., Enum. Filic.: 197, 282. 1824.
Pteris L., Sp. Pl. 2: 1073. 1753. Lectotype (desig- Lectotype (designated by J.Sm., Hist. Fil.: 154.
nated by J.Sm., Hist. Fil.: 295. 1875): Pteris longifolia 1875): Antrophyum plantagineum (Cav.) Kaulf. (
L. Circumscription sensu Zhang et al. (2015). Hemionitis plantaginea Cav.). Circumscription
Includes: Afropteris Alston; Anopteris Prantl ex sensu Schuettpelz et al. (2016). Monophyletic
Diels; Ochropteris J.Sm.; Neurocallis Fe e; Platyzoma (Schuettpelz et al., 2016). Perhaps 40 species.
R.Br. Monophyletic (Chao et al., 2014; Zhang et al.,
2015). Perhaps 250 species. Haplopteris C.Presl, Tent. Pterid.: 141. 1836. Type:
Haplopteris scolopendrina (Bory) C.Presl ( Pteris
Pterozonium Fe e, Me
m. Soc. Hist. Nat. Strasbourg scolopendrina Bory). Circumscription sensu
4(1): 202. 1850. Type: Pterozonium reniforme Schuettpelz et al. (2016). Includes: Pleurofossa
(Mart.) Fee ( Gymnogramma reniformis Mart.). Nakai ex H.Ito; Monogramma Comm. ex Schkuhr
Circumscription sensu Tryon et al. (1990). Mono- (proposal to conserve Haplopteris against Mono-
phyletic (Cochran et al., 2014), but relatively few gramma pending; Chen et al., 2016). Mono-
species sampled. 14 species. phyletic (Schuettpelz et al., 2016). Perhaps 40
species.
Syngramma J.Sm., London J. Bot. 4: 168–169. 1845.
Lectotype (designated by J.Sm., Hist. Fil.: 152. Hecistopteris J.Sm., London J. Bot. 1: 193. 1842.
1875): Syngramma alismifolia (C.Presl) J.Sm. ( Type: Hecistopteris pumila (Spreng.) J.Sm. (
Diplazium alismifolium C.Presl). Circumscription Gymnogramma pumila Spreng.). Circumscription
sensu Tryon et al. (1990). Only one species sensu Schuettpelz et al. (2016). Presumably
sampled to date, but assumed to be monophyletic. monophyletic, but only one species sampled to
About 15 species. date (Schuettpelz et al., 2016). Three species.
Taenitis Willd. ex Schkuhr, Kl. Linn. Pfl.-Syst. 1: 20. Polytaenium Desv., Me m. Soc. Linn. Paris 6: 174,
1804. Type: Taenitis pteroides Willd. ex Schkuhr. 218. 1827. Type: Polytaenium lanceolatum (Sw.)
Circumscription sensu Tryon et al. (1990). Mono- Desv. ( Vittaria lanceolata Sw.). Circumscription
phyletic (Cochran et al., 2014), but sampling sensu Schuettpelz et al. (2016). Includes: Anetium
insufficient. About 15 species. Splitg. Monophyletic (Schuettpelz et al., 2016).
About 10 species.
Tryonia Schuettp., J.Prado & A.T.Cochran, Phyto-
Keys 35: 35–37. 2014. Type: Tryonia myriophylla Radiovittaria (Benedict) E.H.Crane, Syst. Bot. 22:
(Sw.) Schuettp., J.Prado & A.T.Cochran ( Gym- 514–515. 1997 (1998). Type: Radiovittaria remota
nogramma myriophylla Sw.). Circumscription sensu (F
ee) E.H.Crane ( Vittaria remota Fee). Circum-
Cochran et al. (2014). Monophyletic (Cochran et al., scription sensu Schuettpelz et al. (2016). Mono-
2014). Four species. phyletic (Schuettpelz et al., 2016). About 10
species.
Subfamily Vittarioideae Link, Fil. Spec.: 116. 1841.
Circumscription equivalent to “adiantoids“ in Schuett- Rheopteris Alston, Nova Guinea, n.s. 7: 2. 1956.
pelz et al. (2007). Monophyletic (Prado et al., 2007; Type: Rheopteris cheesmaniae Alston. Circumscrip-
Schuettpelz et al., 2007; Rothfels & Schuettpelz, 2014). tion sensu Schuettpelz et al. (2016). Monotypic.
12 genera and an estimated 345 species.
Scoliosorus T.Moore, Index Fil. xxix. 1857. Type:
Adiantum L., Sp. Pl. 2: 1094. 1753. Lectotype Scoliosorus ensiformis (Hook.) T.Moore ( Antro-
(designated by J.Sm., Hist. Fil.: 274. 1875): Adiantum phyum ensiforme Hook.). Circumscription sensu
capillus-veneris L. Circumscription sensu Tryon et al. Schuettpelz et al. (2016). Monotypic (Schuettpelz
(1990). Monophyletic (Rothfels & Schuettpelz, et al., 2016).
2014; Pryer et al., 2016). About 225 species.
Vaginularia Fe e, Me
m. Foug., 3. Hist. Vittar.: 30–31.
Ananthacorus Underw. & Maxon, Contr. U.S. Natl. 1852. Type: Vaginularia trichoidea Fee. Circumscrip-
Herb. 10: 487. 1908. Type: Ananthacorus angustifo- tion sensu Schuettpelz et al. (2016). Includes:
lius (Sw.) Underw. & Maxon ( Pteris angustifolia Diclidopteris Brack. Monophyletic (Schuettpelz
Sw.). Circumscription sensu Schuettpelz et al. et al., 2016). Four species.
(2016). Monotypic (Schuettpelz et al., 2016).
Vittaria Sm., Me m. Acad. Roy. Sci. (Turin) 5: 413,
Antrophyopsis (Benedict) Schuettp., Taxon 65(4): t.9, f.5. 1793. Type: Vittaria lineata (L.) Sm. ( Pteris
717. 2016. Type: Antrophyopsis boryana (Willd.) lineata L.). Circumscription sensu Schuettpelz et al.

PPG I 579
(2016). Monophyletic (Schuettpelz et al., 2016). Herb. 13: 160. 1913): Bommeria ehrenbergiana
Seven species. (Klotzsch) Underw. ( Gymnogramma ehrenbergi-
ana Klotzsch). Circumscription sensu Ranker &
Subfamily Cheilanthoideae Horvat, Acta Bot. Inst. Haufler (1990). Monophyletic based on limited
Bot. Univ. Zagreb. 2: 114. 1927, as “Cheilanthinae”. sampling (two species in Eiserhardt et al., 2011).
Circumscription equivalent to “cheilanthoids” in Five species.
Schuettpelz et al. (2007). Monophyletic (Rothfels &
Schuettpelz, 2014). 23 genera and an estimated 426 Calciphilopteris Yesilyurt & H.Schneid., Phytotaxa
species. 7: 52. 2010. Type: Calciphilopteris ludens (Wall. ex
Hook.) Yesilyurt & H.Schneid. ( Pteris ludens Wall.
Adiantopsis Fe e, Me m. Foug., 5. Gen. Filic.: 145. ex Hook.). Circumscription sensu Yesilyurt &
1852. Lectotype (designated by C.Chr., Index Filic. Schneider (2010). Presumably monophyletic based
e ( Adiantum
xli. 1906): Adiantopsis radiata (L.) Fe on morphological similarity (only one species
radiatum L.). Circumscription sensu Link-Pe rez sampled; Schuettpelz et al., 2007; Eiserhardt
et al. (2011) with additions by Link-Pe rez & Hickey et al., 2011; Rothfels & Schuettpelz, 2014). Four
(2011) and Schuettpelz et al. (2014). Monophyletic species.
(Link-Perez et al., 2011; Schuettpelz et al., 2014).
About 35 species. Cheilanthes Sw., Syn. Fil. 5, 126. 1806. Lectotype
(designated by Maxon, Sci. Surv. Porto Rico &
Aleuritopteris Fe e, Mem. Foug., 5. Gen. Filic.: 153– Virgin Islands 6(3): 428. 1926): Cheilanthes micro-
154. 1852. Lectotype (designated by C.Chr., Index pteris Sw. Circumscription sensu Zhang & Yatskie-
e
Filic. xlii. 1906): Aleuritopteris farinosa (Forssk.) Fe vych (2013) but excluding Allosorus and Mildella.
( Pteris farinosa Forssk.). Circumscription sensu Includes: Cheilosoria Trevis.; Negripteris Pic.Serm.;
Zhang et al. (2013a). Includes: Leptolepidium K.H. Neurosoria Mett. ex Kuhn. Not monophyletic as
Shing & S.K.Wu; Sinopteris C.Chr. & Ching. Not circumscribed (Eiserhardt et al., 2011). About 100
monophyletic as circumscribed (see Eiserhardt species.
et al., 2011). About 40 species.
Cheiloplecton Fee, Mem. Foug., 7. Ic. Esp. Nouv.:
Allosorus Bernh., Neues J. Bot. 1(2): 36. 1805. 33, 135, t.20. 1857. Type: Cheiloplecton rigidum
Lectotype (designated by Pic.Serm., Webbia 9: (Sw.) F ee ( Pteris rigida Sw.). Circumscription
394. 1953): Allosorus pusillus (Willd. ex Bernh.) sensu Mickel & Smith (2004). Monotypic (Rothfels
Bernh. ( Adiantum pusillum Willd. ex Bernh.). et al., 2008; Johnson et al., 2012).
Circumscription sensu Christenhusz (2012). In-
cludes: Oeosporangium De Visiani. Questionably Doryopteris J.Sm. emend. J.C.Yesilyurt, J. Bot.
monophyletic as circumscribed. Eight species. (Hooker) 3: 404. 1841. Lectotype (designated by
C.V.Morton, Amer. Fern J. 34: 26. 1944): Doryop-
Argyrochosma (J.Sm.) Windham, Amer. Fern J. 77 teris palmata (Willd.) J.Sm. ( Pteris palmata
(2): 38. 1987. Lectotype (designated by C.Chr., Willd.). Circumscription sensu Yesilyurt et al.
Index Filic. xl. 1906): Argyrochosma nivea (Poir.) (2015). Includes: Bakeriopteris C.Chr.; Cassebeera
Windham ( Pteris nivea Poir.). Circumscription Kaulf.; Heteropteris F
ee; Tryonella Pic.Serm. Mono-
sensu Windham (1987) with additions by Wang phyletic (Yesilyurt et al., 2015). 21 species.
et al. (2015). Monophyletic (Sigel et al., 2011).
About 20 species. Gaga Pryer, F.W.Li & Windham, Syst. Bot. 37(4):
855. 2012. Type: Gaga marginata (Kunth) F.W.Li &
Aspidotis (Nutt. ex Hook.) Copel., Ann. Cryptog. Windham ( Cheilanthes marginata Kunth). Cir-
Phytopathol. 5 [Gen. Fil.]: 68. 1947. Type: Aspidotis cumscription sensu Li et al. (2012b). Monophyletic
californica (Hook.) Nutt. ex Copel. ( Hypolepis (Li et al., 2012b). 19 species.
californica Hook.). Circumscription sensu Li et al.
(2012b). Monophyletic (Li et al., 2012b). Four Hemionitis L., Sp. Pl. 2: 1077 1753. Lectotype
species. (designated by Kaulf., Enum. Filic.: 68, 198. 1824):
Hemionitis palmata L. Circumscription sensu Tryon
Astrolepis D.M.Benham & Windham, Amer. Fern J. et al. (1990). Includes: Gymnopteris Bernh.; Gym-
82(2): 55. 1992. Type: Astrolepis sinuata (Lag. ex nogramma Desv.; Neurogramma Link. Monophy-
Sw.) D.M.Benham & Windham ( Acrostichum letic (four species sampled in Eiserhardt et al.,
sinuatum Lag. ex Sw.). Circumscription sensu 2011). Five species.
Benham & Windham (1992) with additions by
Beck et al. (2010). Monophyletic (Beck et al., 2010). Lytoneuron (Klotzsch) Yesilyurt, Phytotaxa 221(2):
Eight species. 116. 2015. Type: Lytoneuron lomariaceum (Klotzsch)
Yesilyurt ( Doryopteris lomariacea Klotzsch).
Bommeria E.Fourn., Dict. Bot. 1: 448. 1876 (1877). Circumscription sensu Yesilyurt et al. (2015).
Lectotype (designated by Maxon, Contr. U.S. Natl. Monophyletic (Yesilyurt et al., 2015). 12 species.

Mildella Trevis., Reach Ist. Lombardo Sci., Rendi- Circumscription sensu Schuettpelz et al. (2015).
conti 9: 810. 1877. Type: Mildella intramarginalis Monophyletic (Schuettpelz et al., 2007). Six
(Kaulf. ex Link) Trevis. ( Pteris intramarginalis species.
Kaulf. ex Link). Circumscription sensu Hall &
Lellinger (1967). Not monophyletic as circum- Trachypteris Andre ex Christ, Neue Denkschr. Allg.
scribed; New World species (including type) and Schweiz. Ges. Gesamten Naturwiss. 36: 150. 1899.
Old World species belong to different clades. Type: Trachypteris aureonitens (Hook.) Andr e ex
Seven species. Christ ( Acrostichum aureonitens Hook.). Circum-
scription sensu Ramos-Giacosa et al. (2011).
Myriopteris Fee emend. Grusz & Windham, Me m. Includes: Saffordia Maxon. Monophyly not as-
Foug., 5. Gen. Filic.: 148. 1852. Lectotype (desig- sessed (only one species sampled; Eiserhardt et al.,
nated by Copel., Ann. Cryptog. Phytopathol. 5 2011). Four species.
[Gen. Fil.]: 65. 1947): Myriopteris marsupianthes
e. Circumscription sensu Grusz & Windham
Fe Suborder Dennstaedtiineae Schwartsb. & Hovenkamp,
(2013). Includes: Pomatophytum M.E.Jones. Mono- this classification (see below). Circumscription equivalent
phyletic (Grusz et al., 2014). About 45 species. to family Dennstaedtiaceae in this classification and in
Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer,
Notholaena R.Br., Prodr.: 145. 1810. Lectotype 2007). An estimated 265 species.
(designated by J.Sm., Hist. Fil.: 278. 1875): Notho-
laena trichomanoides (L.) Desv. ( Pteris tricho- 31. Family Dennstaedtiaceae Lotsy, Vort. Bot. Stammes-
manoides L.). Circumscription sensu Rothfels et al. gesch. 2: 655. 1909. Circumscription sensu Smith et al.
(2008) with additions by Yatskievych & Arbelaez (2006b). Includes: Hypolepidaceae Pic.Serm.; Mona-
(2008). Includes: Chrysochosma (J.Sm.) K€ ummerle. chosoraceae Ching; Pteridiaceae Ching. Monophyletic
See Yatskievych & Smith (2003) for details (Schuettpelz & Pryer, 2007). Ten genera and an
regarding arguments over the typification of estimated 265 species.
Notholaena. Not monophyletic as circumscribed
(Rothfels et al., 2008; Johnson et al., 2012). About Blotiella R.M.Tryon, Contr. Gray Herb. 191: 96. 1962.
30 species. Type: Blotiella glabra (Bory) R.M.Tryon ( Lonchitis
glabra Bory). Circumscription sensu Tryon (1962)
Ormopteris J.Sm. ex J.Sm., Hist. Fil.: 281. 1875. and Kramer (1990d). Monophyletic (Perrie et al.,
Type: Ormopteris gleichenioides (Gardner) J.Sm. ( 2015), although only two species were sampled.
Cassebeera gleichenioides Gardner). Circumscrip- Perhaps 20 species.
tion sensu Yesilyurt et al. (2015). Monophyletic
(Yesilyurt et al., 2015). Six species. Dennstaedtia Bernh., J. Bot. (Schrader) 1800(2):
124. 1801. Type: Dennstaedtia flaccida (G.Forst.)
Paragymnopteris K.H.Shing, Indian Fern J. 10: 227. Bernh. ( Trichomanes flaccidum G.Forst.). Circum-
1993. Type: Paragymnopteris marantae (L.) K.H. scription sensu Kramer (1990d). Includes: Adectum
Shing ( Acrostichum marantae L.). Circumscrip- Link; Coptodipteris Nakai & Momose; Costaricia
tion sensu Zhang & Ranker (2013b). Not monophy- Christ; Emodiopteris Ching & S.K.Wu; Fuziifilix Kanai
letic as circumscribed. Five species. & Momose; Paradennstaedtia Tagawa; Patania C.
Presl; Sitobolium Desv. Not monophyletic as
Parahemionitis Panigrahi, Amer. Fern J. 83(3): 90, circumscribed; needs further sampling to include
f.1. 1993. Type: Parahemionitis arifolia (Burm.f.) several competing type species (Perrie et al., 2015).
Panigrahi ( Asplenium arifolium Burm.f.). Circum- About 70 species.
scription sensu Zhang & Ranker (2013a). Mono-
typic. Histiopteris (J.Agardh) J.Sm., Hist. Fil.: 294: 1875.
Type: Histiopteris vespertilionis (Labill.) J.Sm. (
Pellaea Link, Fil. Spec.: 59. 1841. Lectotype Pteris vespertilionis Labill.). Circumscription sensu
(designated by C.Chr., Index Filic. xxxix. 1906): Kramer (1990d). Includes: Lepidocaulon Copel.
Pellaea atropurpurea (L.) Link ( Pteris atropur- Monophyletic (Perrie et al., 2015), although
purea L.). Circumscription sensu Tryon et al. (1990) only two species were sampled. About seven
but excluding Ormopteris. Includes: Choristosoria species.
Mett. ex Kuhn; Paraceterach Copel.; Pellaeopsis J.
Sm.; Platyloma J.Sm.; Pteridella Kuhn. Not mono- Hypolepis Bernh., Neues J. Bot. 1(2): 34. 1805. Type:
phyletic as circumscribed (Yesilyurt et al., 2015). Hypolepis tenuifolia (G.Forst.) Bernh. ex C.Presl (
About 40 species. Lonchitis tenuifolia G.Forst.). Circumscription sensu
Kramer (1990d) and Schwartsburd & Prado (2015,
Pentagramma Yatsk., Windham & E.Wollenw., 2016). Mostly monophyletic (Wolf, 1995; Perrie
Amer. Fern J. 80(1): 13. 1990. Type: Pentagramma et al., 2015), but a few species may need to be
triangularis (Kaulf.) Yatsk., Windham & E. transferred to a different genus (Brownsey, 1983).
Wollenw. ( Gymnogramma triangularis Kaulf.). Perhaps 80 species.

PPG I 581
Leptolepia Mett. ex Prantl, Arbeiten K€ onigl. Bot. sensu Rothfels et al. (2012b). Monophyletic
Gart. Breslau 1: 23. 1892. Lectotype (designated by (Rothfels et al., 2012a, 2013). At least three species.
C.Chr., Index Filic. xxviii. 1906): Leptolepia novae-
zelandiae (Colenso) Mett. ex Diels ( Davallia Cystopteris Bernh., Neues J. Bot. 1(2): 26. 1805.
novae-zelandiae Colenso). Circumscription sensu Lectotype (designated by J.Sm., Hist. Fil.: 236.
Kramer (1990d). Monotypic. 1875): Cystopteris fragilis (L.) Bernh. ( Polypodium
fragile L.). Circumscription sensu Rothfels et al.
Microlepia C.Presl, Tent. Pterid.: 124, pl. 4, f. 21–23. (2013) and Wei & Zhang (2014). Includes: Cystoa-
1836. Lectotype (designated by J.Sm., Hist. Fil.: thyrium Ching; Rhizomatopteris A.P.Khokhr. Mono-
260. 1875): Microlepia polypodioides (Sw.) C.Presl phyletic (Rothfels et al., 2012a, 2013; Wei & Zhang,
( Dicksonia polypodioides Sw.). Circumscription 2014). At least 26 species.
sensu Kramer (1990d). Includes: Scypholepia J.Sm.
Monophyletic; sister to the old world clade Gymnocarpium Newman, Phytologist 4: 371. 1851.
of Dennstaedtia (Perrie et al., 2015). About 60 Lectotype (designated by Ching, Contr. Biol. Lab.
species. Sci. Soc. China, Bot. Ser. 9: 38. 1933): Gymno-
carpium dryopteris (L.) Newman ( Polypodium
Monachosorum Kunze, Bot. Zeitung (Berlin) 6: 119. dryopteris L.). Circumscription sensu Sarvela (1978)
1848. Type: Monachosorum davallioides Kunze. and Rothfels et al. (2013). Includes: Currania Copel.
Circumscription sensu Kramer (1990m). Includes: Monophyletic (Rothfels et al., 2012a, 2013). At least
Ptilopteris Hance; Monachosorella Hayata. Mono- eight species.
phyletic (Perrie et al., 2015; Ebihara et al., 2016).
About six species. 33. Family Rhachidosoraceae X.C.Zhang, Phytotaxa 19: 16.
2011. Circumscription sensu Christenhusz et al. (2011)
Oenotrichia Copel., Univ. Calif. Publ. Bot. 16: 82. and Rothfels et al. (2012b). Family consists of a single
1929. Type: Oenotrichia maxima (E.Fourn.) Copel. genus and an estimated eight species.
( Leucostegia maxima E.Fourn.). Circumscription
sensu Kramer (1990d). Monophyletic (Perrie et al., Rhachidosorus Ching, Acta Phytotax. Sin. 9(1): 73.
2015). About two species. 1964. Type: Rhachidorous mesosorus (Makino)
Ching ( Asplenium mesosorum Makino). Circum-
Paesia A.St.-Hil., Voy. Distr. Diam. 1: 381. 1833. Type: scription sensu Lindsay et al. (2012). Likely
Paesia viscosa A.St.-Hil. Circumscription sensu monophyletic (Li et al., 2011; Rothfels et al.,
Kramer (1990d). Monophyletic (Perrie et al., 2012a). Eight species.
2015), although only two species were sampled.
About 15 species. 34. Family Diplaziopsidaceae X.C.Zhang & Christenh.,
Phytotaxa 19: 15. 2011. Circumscription sensu Rothfels
Pteridium Gled. ex Scop., Fl. Carniol.: 169. 1760. et al. (2012b). Likely monophyletic (Kuo et al., 2011;
Type: Pteridium aquilinum (L.) Kuhn ( Pteris Rothfels et al., 2012a). Two genera and an estimated
aquilina L.). Circumscription sensu Kramer (1990d), four species.
Der et al. (2009), and Thomson (2012). Includes:
Cincinalis Gled.; Eupteris Newman; Ornithopteris (J. Diplaziopsis C.Chr., Index Filic. 227. 1905. Type:
Agardh) J.Sm.; Filix-foemina Farwell. Monophyletic Diplaziopsis brunoniana (Wall.) W.M.Chu ( Allan-
(Der et al., 2009; Zhou et al., 2014; Wolf et al., 2015). todia brunoniana Wall.). Circumscription sensu He
Species number uncertain; about 20 morphotypes & Kato (2013). Likely monophyletic (Li et al., 2011;
have been recognized, but molecular evidence Wei et al., 2010; Rothfels et al., 2012a). Three
points to substantially fewer species. Four species. species.
Suborder Aspleniineae H.Schneid. & C.J.Rothf., this Homalosorus Small ex Pic.Serm., Webbia 31(1): 246.
classification (see below). Circumscription equivalent to 1977. Type: Homalosorus pynocarpos (Spreng.) Pic.
“eupolypods II” in Smith et al. (2006b). Monophyletic Serm. ( Asplenium pycnocarpum Spreng.). Cir-
(Schuettpelz & Pryer, 2007; Rothfels et al., 2012a). 11 cumscription sensu Pichi Sermolli (1977a). Mono-
families, 72 genera, and an estimated 2775 species. typic.
32. Family Cystopteridaceae Shmakov, Turczaninowia 4: 35. Family Desmophlebiaceae Mynssen, A.Vasco, Syl-
60. 2001. Circumscription sensu Rothfels et al. (2012b). vestre, R.C.Moran & Rouhan, Taxon 65(1): 19. 2016.
Monophyletic (Rothfels et al., 2012a, 2013; Wei & Circumscription sensu Mynssen et al. (2016). Family
Zhang, 2014). Three genera and an estimated 37 consists of a single genus and two species.
species.
Desmophlebium Mynssen, A.Vasco, Sylvestre, R.C.
Acystopteris Nakai, Bot. Mag. (Tokyo) 47: 180. Moran & Rouhan, Taxon 65(1): 27, f.5. 2016. Type:
1933. Type: Acystopteris japonica (Luerss.) Nakai ( Desmophlebium lechleri (Mett.) Mynssen, A.Vasco,
Cystopteris japonica Luerss.). Circumscription Sylvestre, R.C.Moran & Rouhan ( Asplenium

lechleri Mett.). Circumscription sensu Mynssen Matteuccia Tod., Giorn. Sci. Nat. Econ. Palermo 1
et al. (2016). Only one species included in a (3–4): 235. 1866. Type: Matteuccia struthiopteris
phylogenetic analysis, but assumed to be mono- (L.) Tod. ( Osmunda struthiopteris L.). Circum-
phyletic based on morphological evidence (Myns- scription sensu Gastony & Ungerer (1997) and
sen et al., 2016). Two species. Rothfels et al. (2012b). Monotypic, although
American and European populations could be
36. Family Hemidictyaceae Christenh. & H.Schneid., Phy- treated as distinct species (Koenemann et al.,
totaxa 28: 51. 2011. Circumscription sensu Rothfels 2011).
et al. (2012b) and Christenhusz & Schneider (2011).
Family consists of a single monotypic genus. Onoclea L., Sp. Pl. 2: 1062. 1753. Type: Onoclea
sensibilis L. Circumscription sensu Gastony &
Hemidictyum C.Presl, Tent. Pterid.: 110, t.3, f.24–26. Ungerer (1997) and Rothfels et al. (2012b).
1836. Lectotype (designated by J.Sm., Hist. Fil.: Includes: Pterinodes Siegesb. ex Kuntze; Calypte-
335. 1875): Hemidictyum marginatum (L.) C.Presl ( rium Bernh.; Ragiopteris C.Presl; Riedlea Mirb.
Asplenium marginatum L.). Circumscription sensu Monotypic (Gastony & Ungerer, 1997).
Kramer et al. (1990). Monotypic.
Onocleopsis F.Ballard, Amer. Fern J. 35: 1. 1945.
37. Family Aspleniaceae Newman, Hist. Brit. Ferns: 6. Type: Onocleopsis hintonii F.Ballard. Circumscrip-
1840. Circumscription sensu Smith et al. (2006b) and tion sensu Gastony & Ungerer (1997) and Rothfels
Rothfels et al. (2012b). Monophyletic (Schneider et al., et al. (2012b). Monotypic.
2004a; Schneider et al., 2005). Two genera and an
estimated 730 species. Pentarhizidium Hayata, Bot. Mag. (Tokyo) 42: 345.
1928. Type: Pentarhizidium japonicum Hayata.
Asplenium L., Sp. Pl. 2: 1078. 1753. Lectotype Circumscription sensu Gastony & Ungerer (1997)
(designated by J.Sm., Hist. Fil.: 316. 1875): Asple- and Rothfels et al. (2012b). Monophyletic (Gastony
nium marinum L. Circumscription sensu Schneider & Ungerer, 1997; Rothfels et al., 2012a). The name
et al. (2004a). Includes: Antigramma C.Presl; Pentarhizidium was first used in 1927, in the
Camptosorus Link; Ceterach Willd.; Ceterachopsis context of describing the morphological distinc-
(J.Sm.) Ching; Diellia Brack.; Diplora Baker; Hol- tiveness of the P. orientale stele (Hayata, 1927).
odictyum Maxon; Loxoscaphe T.Moore; Phyllitis However, the paper lacked a description of the
Hill; Pleurosorus F
ee; Schaffneria F
ee ex T.Moore; genus itself and we consider Pentarhizidium and P.
Scolopendrium Adans.; Sinephropteris Mickel. orientale as nomina nuda until the genus was
Monophyletic (Schneider et al., 2004a). At least formally described and a type (P. japonicum) was
700 species. designated (Hayata, 1928). Two species.
Hymenasplenium Hayata, Bot. Mag. (Tokyo) 41 40. Family Blechnaceae Newman, Hist. Brit. Ferns (ed. 2):
(492): 712. 1927. Type: Hymenasplenium unilaterale 8. 1844. Circumscription sensu Smith et al. (2006b).
(Lam.) Hayata ( Asplenium unilaterale Lam.). Monophyletic (Schuettpelz & Pryer 2007; Rothfels
Circumscription sensu Schneider et al. (2004a). et al., 2012a; Gasper et al., 2016a). Three subfamilies, 24
Monophyletic (Schneider et al., 2004a). At least 30 genera, and an estimated 265 species.
species.
Subfamily Stenochlaenoideae (Ching) J.P.Roux, Con-
38. Family Woodsiaceae Herter, Revista Sudamer. Bot. 9: spect. South. Afr. Pteridophyta: 156. 2001. Circumscrip-
14. 1949. Circumscription sensu Christenhusz et al. tion sensu Roux (2001) emend. Gasper et al. (2016b).
(2011) and Rothfels et al. (2012b). Family consists of a Monophyletic (Gasper et al., 2016a). Three genera and
single genus and an estimated 39 species. an estimated 12 species.
Woodsia R.Br., Prodr.: 158, Obs. 4. 1810. Lectotype Salpichlaena J.Sm., Gen. Fil. pl. 93. 1842. Type:
(designated by J.Sm., Hist. Fil.: 237. 1875): Woodsia Salpichlaena volubilis (Kaulf.) J.Sm. ( Blechnum
ilvensis (L.) R.Br. ( Acrostichum ilvense L.). volubile Kaulf.). Circumscription sensu Perrie et al.
Circumscription sensu Shao et al. (2015). Includes: (2014) and Gasper et al. (2016b). Monophyletic
Eriosoriopsis (Kitag.) Ching & S.H.Wu; Cheilanthop- (Gasper et al., 2016a). Three species.
sis Hieron.; Hymenocystis C.A.Mey; Physematium
Kaulf.; Protowoodsia Ching; Woodsiopsis Shmakov. Stenochlaena J.Sm., J. Bot. (Hooker) 3: 401. 1841.
Monophyletic (Rothfels et al., 2012a; Shao et al., Lectotype (designated by Pfeiffer, Nom. 2: 1274.
2015; Shmakov, 2015). 39 species. 1874): Stenochlaena palustris (Burm.f.) Bedd. (
Polypodium palustre Burm.f.). Circumscription
39. Family Onocleaceae Pic.Serm., Webbia 24: 708. 1970. sensu Perrie et al. (2014) and Gasper et al.
Circumscription sensu Smith et al. (2006b). Monophy- (2016b). Monophyletic (Gasper et al., 2016a). We
letic (Gastony & Ungerer, 1997; Rothfels et al., 2012a). follow Copeland (1947) in identifying the name
Four genera and an estimated five species. Acrostichum scandens L. (referred to in both the

PPG I 583
description of the genus and its lectotypification) Blechnum orientale L.). Circumscription sensu
with Stenochlaena palustris (Burm.f.) Bedd. Seven Gasper et al. (2016b). Monophyletic (Gasper
species. et al., 2016a). Two species.
Telmatoblechnum Perrie, D.J.Ohlsen & Brownsey, Blechnum L., Sp. Pl. 2: 1077. 1753. Lectotype
Taxon 63(4). 755. 2014. Type: Telmatoblechnum (designated by J.Sm., Hist. Fil.: 300. 1875):
serrulatum (Rich.) Perrie, D.J.Ohlsen & Brownsey Blechnum occidentale L. Circumscription sensu
( Blechnum serrulatum Rich.). Circumscription Gasper et al. (2016b). Monophyletic (Gasper
sensu Perrie et al. (2014) and Gasper et al. (2016b). et al., 2016a). 30 species.
Monophyletic (Perrie et al., 2014; Gasper et al.,
2016a). Two species. Brainea J.Sm., Cat. Kew Ferns: 5. 1856. Type:
Brainea insignis (Hook.) J.Sm. ( Bowringia insignis
Subfamily Woodwardioideae Gasper, V.A.O.Dittrich & Hook.). Circumscription sensu Gasper et al.
Salino, Phytotaxa 275(3): 197. 2016. Circumscription (2016b). Monotypic (Gasper et al., 2016a).
sensu Gasper et al. (2016b). Monophyletic (Cranfill &
Kato, 2003; Perrie et al., 2014; Gasper et al., 2016a). Cleistoblechnum Gasper & Salino, Phytotaxa 275
Three genera and an estimated 15 species. (3): 207. 2016. Type: Cleistoblechnum eburneum
(Christ) Gasper & Salino ( Blechnum eburneum
Anchistea C.Presl, Epimel. Bot.: 71. 1851. Type: Christ). Circumscription sensu Gasper et al.
Anchistea virginica (L.) C.Presl ( Blechnum virgin- (2016b). Monotypic (Gasper et al., 2016a).
icum L.). Circumscription sensu Gasper et al.
(2016b). Monotypic (Gasper et al., 2016a). Cranfillia Gasper & V.A.O.Dittrich, Phytotaxa 275
(3): 207. 2016. Type: Cranfillia fluviatilis (R.Br.)
Lorinseria C.Presl, Epimel. Bot.: 72. 1851. Lectotype Gasper & V.A.O.Dittrich ( Stegania fluviatilis
(designated by J.Sm., Hist. Fil.: 310. 1875): Lor- R.Br.). Circumscription sensu Gasper et al.
inseria areolata (L.) C.Presl ( Acrostichum areola- (2016b). Monophyletic (Gasper et al., 2016a). 11
tum L.). Circumscription sensu Gasper et al. species.
(2016b). Lorinseria C.Presl appears to be a later
homonym of Lorinsera Opiz, and will therefore Diploblechnum Hayata, Bot. Mag. (Tokyo) 41. 702.
most likely require a new name (or conservation). 1927. Type: Diploblechnum integripinnulum Hayata.
Monotypic (Gasper et al., 2016a). Circumscription sensu Gasper et al. (2016b).
Includes: Pteridoblechnum Hennipman and Steeni-
Woodwardia Sm., M em. Acad. Roy. Sci. Turin 5. 411. sioblechnum Hennipman. Monophyletic (Gasper
1793. Lectotype (designated by J.Sm., Hist. Fil.: 310. et al., 2016a). Six species.
1875): Woodwardia radicans (L.) Sm. ( Blechnum
radicans L.). Circumscription sensu Gasper et al. Doodia R.Br., Prodr. Fl. Nov. Holl.: 151. 1810.
(2016b). Includes: Chieniopteris Ching. Monophy- Lectotype (designated by J.Sm., Hist. Fil.: 309.
letic (Gasper et al., 2016a). 13 species. 1875): Doodia aspera R.Br. Circumscription sensu
Gasper et al. (2016b). Monophyletic (Gasper et al.,
Subfamily Blechnoideae Gasper, V.A.O.Dittrich & 2016a). 19 species.
Salino, Phytotaxa 275(3): 201. 2016. Circumscrip-
tion sensu Gasper et al. (2016b). Monophyletic (Gasper Icarus Gasper & Salino, Phytotaxa 275(3): 209.
et al., 2016a). 18 genera and an estimated 238 2016. Type: Icarus filiformis (A.Cunn.) Gasper &
species. Salino ( Lomaria filiformis A.Cunn.). Circumscrip-
tion sensu Gasper et al. (2016b). Monotypic
Austroblechnum Gasper & V.A.O.Dittrich, Phyto- (Gasper et al., 2016a).
taxa 275(3): 202. 2016. Type: Austroblechnum
penna-marina (Poir.) Gasper & V.A.O.Dittrich ( Lomaria Willd., Mag. Neuesten Entdeck. Ge-
Polypodium penna-marina Poir.). Circumscription sammten Naturk. Ges. Naturf. Freunde Berlin 3:
sensu Gasper et al. (2016b). Monophyletic (Gasper 160. 1809. Lectotype (designated by J.Sm., Hist.
et al., 2016a). 40 species. Fil.: 303. 1875): Lomaria nuda (Labill.) Willd. (
Onoclea nuda Labill.). Circumscription sensu
Blechnidium T.Moore, Index Fil. clv. 1860; Brit. Gasper et al. (2016b). Includes: Stegania R.Br.
Ferns: 210. 1860. Type: Blechnidium melanopus Monophyletic (Gasper et al., 2016a). Six species.
(Hook.) T.Moore ( Blechnum melanopus Hook.).
Circumscription sensu Gasper et al. (2016b). Lomaridium C.Presl, Epimel. Bot.: 154. 1851. Lecto-
Monotypic (Gasper et al., 2016a). type (designated by C.Chr., Index Filic. xxxv. 1906):
Lomaridium plumieri (Desv.) C.Presl ( Lomaria
Blechnopsis C.Presl, Epimel. Bot.: 115. 1851. Lecto- plumieri Desv.). Circumscription sensu Gasper et al.
type (designated by Pic.Serm., Webbia 28: 456. (2016b). Monophyletic (Gasper et al., 2016a). 16
1973): Blechnopsis orientalis (L.) C.Presl ( species.

Lomariocycas (J.Sm.) Gasper & A.R.Sm., Phytotaxa Diplazium Sw., J. Bot. (Schrader) 1800(2): 4, 61.
275(3): 212. 2016. Type: Lomaria boryana (Sw.) 1801. Lectotype (designated by J.Sm., Hist. Fil.: 325.
Willd. ( Onoclea boryana Sw.). Circumscription 1875): Diplazium plantaginifolium (L.) Urb. (
sensu Gasper et al. (2016b). Monophyletic (Gasper Asplenium plantaginifolium L.). Circumscription
et al., 2016a). 19 species. sensu Wei et al. (2013). Includes: Allantodia R.Br.;
Anisogonium C.Presl; Callipteris Bory; Monomelan-
Neoblechnum Gasper & V.A.O.Dittrich, Phytotaxa gium Hayata. Monophyletic (Wei et al., 2013, 2015).
275(3): 214. 2016. Type: Neoblechnum brasiliense About 350 species.
(Desv.) Gasper & V.A.O.Dittrich ( Blechnum
brasiliense Desv.). Circumscription sensu Gasper 42. Family Thelypteridaceae Ching ex Pic.Serm., Webbia
et al. (2016b). Monotypic (Gasper et al., 2016a). 24: 709. 1970. Circumscription sensu Smith & Cranfill
(2002), He & Zhang (2012), and Almeida et al. (2016).
Oceaniopteris Gasper & Salino, Phytotaxa 275(3): Monophyletic (Smith & Cranfill, 2002; Rothfels
214. 2016. Type: Oceaniopteris gibba (Labill.) Gasper et al., 2012a; Almeida et al., 2016). Two subfamilies,
& Salino ( Lomaria gibba Labill.). Circumscription 30 genera, and an estimated 1034 species.
sensu Gasper et al. (2016b). Monophyletic (Gasper
et al., 2016a). Eight species. Subfamily Phegopteridoideae Salino, A.R.Sm. &
T.E.Almeida, this classification (see below). Circum-
Parablechnum C.Presl, Epimel. Bot.: 109. 1851. scription sensu Smith & Cranfill (2002), He & Zhang
Lectotype (designated by Fe e, Me
m. Foug., 5. (2012), and Almeida et al. (2016). Monophyletic (Smith
Gen. Filic.: 82. 1852): Parablechnum procerum (G. & Cranfill, 2002; He & Zhang, 2012; Rothfels et al.,
Forst.) C.Presl ( Osmunda procera G.Forst.). 2012a; Almeida et al., 2016). Three genera and an
Circumscription sensu Gasper et al. (2016b). estimated 34 species.
Monophyletic (Gasper et al., 2016a). 65 species.
Macrothelypteris (H.Ito) Ching, Acta Phytotax. Sin.
Sadleria Kaulf., Enum. Filic.: 161. 1824. Type: 8(4): 308. 1963. Type: Macrothelypteris oligophle-
Sadleria cyatheoides Kaulf. Circumscription sensu bia (Baker) Ching ( Nephrodium oligophlebium
Perrie et al. (2014) and Gasper et al. (2016b). Baker). Circumscription sensu Almeida et al.
Monophyletic (Gasper et al., 2016a). Six species. (2016). Monophyletic (Smith & Cranfill, 2002;
He & Zhang, 2012; Almeida et al., 2016). 10
Struthiopteris Scop., Fl. Carn.: 168. 1760. Lectotype species.
(designated by Weis, Pl. Crypt. Fl. Gott.: 286. 1770):
Struthiopteris spicant (L.) Weis ( Lomaria spicant Phegopteris (C.Presl) F ee, M
em. Foug., 5. Gen.
L.). Circumscription sensu Gasper et al. (2016b). Filic.: 242. 1852. Lectotype (designated by Ching,
Includes: Spicantopsis Nakai. Monophyletic Acta Phytotax. Sin. 8(4): 312. 1963): Phegopteris
(Gasper et al., 2016a). Five species. polypodioides F ee ( Polypodium phegopteris L.).
Circumscription sensu Almeida et al. (2016).
41. Family Athyriaceae Alston, Taxon 5: 25. 1956. Circum- Monophyletic (Smith & Cranfill, 2002; He & Zhang,
scription sensu Rothfels et al. (2012b). Monophyletic 2012; Almeida et al., 2016). For discussion of the
(Rothfels et al., 2012a). Three genera and an estimated authority and typification, see McNeill and Pryer
650 species. (1985). Four species.
Athyrium Roth, Tent. Fl. Germ. 3(1): 31. 1800. Pseudophegopteris Ching, Acta Phytotax. Sin. 8(4):
Lectotype (designated by J.Sm., Hist. Fil.: 327. 313. 1963. Type: Pseudophegopteris pyrrhorachis
1875): Athyrium filix-femina (L.) Roth ( Polypo- (Kunze) Ching ( Polypodium pyrrhorachis Kunze).
dium filix-femina L.). Circumscription equivalent to Circumscription sensu Almeida et al. (2016).
“athyriids“ in Rothfels et al. (2012b). Includes: Monophyletic (Smith & Cranfill, 2002; He & Zhang,
Anisocampium C.Presl; Kuniwatsukia Pic.Serm.; 2012; Almeida et al., 2016). 20 species.
Pseudocystopteris Ching; Neoathyrium Ching &
Z.R.Wang; Cornoathyrium Nakaike; Cornopteris Subfamily Thelypteridoideae C.F.Reed, Phytologia 17:
Nakai. Monophyletic (Adjie et al., 2008; Rothfels 254. 1968. Circumscription sensu Smith & Cranfill
et al., 2012a). About 230 species. (2002), He & Zhang (2012), and Almeida et al. (2016).
Monophyletic (Smith & Cranfill, 2002; He & Zhang,
Deparia Hook. & Grev., Icon. Filic. 2(8): pl.154. 1830. 2012; Almeida et al., 2016). 27 genera and an estimated
Type: Deparia macraei Hook. & Grev. Circumscrip- 1000 species.
tion sensu Kuo et al. (2016b). Includes: Depazium
Nakaike; Lunathyrium Koidz.; Dryoathyrium Amauropelta Kunze, Farrnkr€auter 1: 86. 1843.
Ching; Parathyrium Holttum; Athyriopsis Ching; Type: Amauropelta breutelii Kunze. Circumscrip-
Dictyodroma Ching; Triblemma (J.Sm.) Ching. tion sensu Almeida et al. (2016). Monophyletic
Monophyletic (Kuo et al., 2016b). About 70 (Smith & Cranfill, 2002; Almeida et al., 2016). About
species. 215 species.

PPG I 585
Amblovenatum J.P.Roux, Strelitzia 23: 200. 2009. Meniscium Schreb., Gen. Pl. ed. 8(a) 2: 757. 1791.
Replacement name for Amphineuron Holttum non Type: Meniscium reticulatum (L.) Sw. ( Polypodium
Amphineurion (A.DC.) Pichon. Type: Amblovena- reticulatum L.). Circumscription sensu Almeida
tum opulentum (Kaulf.) J.P.Roux ( Aspidium et al. (2016). Monophyletic (Smith & Cranfill,
opulentum Kaulf.). Circumscription sensu Almeida 2002; He & Zhang, 2012; Almeida et al., 2016). 27
et al. (2016). Monophyletic (He & Zhang, 2012; species.
Almeida et al., 2016). 12 species.
Menisorus Alston, Bol. Soc. Brot., s
er. 2. 30: 20.
Ampelopteris Kunze, Bot. Zeitung (Berlin) 6: 114. 1956. Type: Menisorus pauciflorus (Hook.) Alston
1848. Type: Ampelopteris elegans Kunze. Circum- ( Meniscium pauciflorum Hook.). Circumscrip-
scription sensu Almeida et al. (2016). Monotypic tion sensu Holttum (1974a). Not yet included in a
(He & Zhang, 2012; Almeida et al., 2016). phylogenetic study. Monotypic.
Chingia Holttum, Blumea 19(1): 31. 1971. Type: Mesophlebion Holttum, Blumea 19(1): 29. 1971.
Chingia ferox (Blume) Holttum ( Aspidium ferox Type: Mesophlebion crassifolium (Blume) Holttum
Blume). Circumscription sensu Almeida et al. ( Aspidium crassifolium Blume). Circumscription
(2016). Monophyletic (Smith & Cranfill, 2002; sensu Holttum (1975). Monophyly uncertain (only
He & Zhang, 2012; Almeida et al., 2016). 20 one species sampled; Almeida et al., 2016). 15
species. species.
v., Fl. Kouy-Tche

Christella H.Le ou 472. 1915. Lecto- Mesopteris Ching, Acta Phytotax. Sin. 16(4): 21.
type (designated by Holttum, Taxon 20: 534. 1971): 1978. Type: Mesopteris tonkinensis (C.Chr.) Ching
Christella parasitica (L.) H.Lev. ( Polypodium ( Dryopteris tonkinensis C.Chr.). Circumscription
parasiticum L.). Circumscription sensu Almeida sensu Lin et al. (2013). Monotypic.
et al. (2016). Not monophyletic as circumscribed
(Almeida et al., 2016). Perhaps 70 species. Metathelypteris (H.Ito) Ching, Acta Phytotax.
Sin. 8(4): 305. 1963. Type: Metathelypteris
Coryphopteris Holttum, Blumea 19(1): 33. 1971. gracilescens (Blume) Ching ( Aspidium graciles-
Type: Coryphopteris viscosa (Baker) Holttum ( cens Blume). Circumscription sensu Almeida et al.
Nephrodium viscosum Baker). Circumscription (2016). Monophyletic (Smith & Cranfill, 2002;
sensu Almeida et al. (2016). Includes: Parathelyp- He & Zhang, 2012; Almeida et al., 2016). 12
teris Ching (pro parte). Monophyletic (Smith & species.
Cranfill, 2002; Almeida et al., 2016). 47 species.
Nannothelypteris Holttum, Blumea 19(1): 38. 1971.
Cyclogramma Tagawa, Acta Phytotax. Geobot. 7 Type: Nannothelypteris aoristisora (Harr.) Holt-
(1): 52. 1938. Type: Cyclogramma simulans (Ching) tum ( Polypodium aoristisorum Harr.). Circum-
Tagawa ( Thelypteris simulans Ching). Circum- scription sensu Holttum (1971). Monophyly
scription sensu Almeida et al. (2016). Presumably uncertain (only one species sampled; Almeida
monophyletic (He & Zhang, 2012; Almeida et al., et al., 2016). Three species.
2016). Eight species.
Oreopteris Holub, Folia Geobot. Phytotax. 4
Cyclosorus Link, Hort. Berol. 2: 128. 1833. Type: (1): 46. 1969. Type: Oreopteris limbosperma
Cyclosorus gongylodes (Schkuhr) Link ( Aspidium (All.) Holub ( Polypodium limbospermum All.).
gongylodes Schkuhr). Circumscription sensu Al- Circumscription sensu Almeida et al. (2016).
meida et al. (2016). Monophyletic (Almeida et al., Monophyletic (Smith & Cranfill, 2002; He
2016). Two species. & Zhang, 2012; Almeida et al., 2016). Three
species.
Glaphyropteridopsis Ching, Acta Phytotax. Sin. 8
(4): 320. 1963. Type: Glaphyropteridopsis erubes- Parathelypteris (H.Ito) Ching, Acta Phytotax. Sin.
cens (Wall. ex Hook.) Ching ( Polypodium 8: 300. 1963. Type: Parathelypteris glanduligera
erubescens Wall. ex Hook.). Circumscription sensu (Kunze) Ching ( Aspidium glanduligerum
Holttum (1971). Presumably monophyletic (Al- Kunze). Circumscription sensu Holttum (1971).
meida et al., 2016). Four species. Not monophyletic as currently circumscribed
(Almeida et al., 2016). About 15 species.
Goniopteris C.Presl, Tent. Pterid.: 181, pl.7, f.9–11.
1836. Lectotype (designated by J.Sm., Hist. Fil.: 191. Plesioneuron (Holttum) Holttum, Blumea 22: 232.
1875): Goniopteris crenata (Sw.) C.Presl ( Poly- 1975. Type: Plesioneuron tuberculatum (Ces.)
podium crenatum Sw.). Circumscription sensu Holttum ( Nephrodium tuberculatum Ces.).
Almeida et al. (2016). Monophyletic (Smith & Circumscription sensu Holttum (1975). Mono-
Cranfill, 2002; He & Zhang, 2012; Almeida et al., phyly uncertain (only one species sampled;
2016). About 120 species. Almeida et al., 2016). 30 species.

Pneumatopteris Nakai, Bot. Mag. (Tokyo) 47: Suborder Polypodiineae Dumort., Anal. Fam. Pl.: 67. 1829.
179. 1933. Type: Pneumatopteris callosa (Blume) Circumscription equivalent to “eupolypods I“ in Smith
Nakai ( Aspidium callosum Blume). Circum- et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007;
scription sensu Holttum (1971). Monophyly un- Rothfels et al., 2015). Nine families, 108 genera, and an
certain (only two species sampled; He & estimated 4208 species.
Zhang, 2012; Almeida et al., 2016). About 80
species. 43. Family Didymochlaenaceae Ching ex Li Bing Zhang &
Liang Zhang, Taxon 64(1): 34. 2015. Circumscription
Pronephrium C.Presl, Epimel. Bot.: 258. 1851. sensu Zhang & Zhang (2015). Family consists of a single
Lectotype (designated by Holttum, Novit. Bot. monotypic genus.
1968: 48): Pronephrium lineatum (Blume) C.Presl
( Aspidium lineatum Blume). Circumscription Didymochlaena Desv., Mag. Neuesten Entdeck.
sensu Holttum (1971). Includes: Abacopteris Fee; Gesammten Naturk. Ges. Naturf. Freunde Berlin 5:
Dimorphopteris Tagawa & K.Iwats. Not mono- 303. 1811. Type: Didymochlaena sinuosa Desv.
phyletic as circumscribed (Almeida et al., 2016). Circumscription sensu Zhang & Zhang (2015).
68 species. Monotypic, although undescribed species may
exist.
Pseudocyclosorus Ching, Acta Phytotax. Sin. 8(4):
322. 1963. Type: Pseudocyclosorus tylodes (Kunze) 44. Family Hypodematiaceae Ching, Acta Phytotax. Sin. 13
Ching ( Aspidium tylodes Kunze). Circumscrip- (1): 96. 1975. Circumscription sensu Zhang & Zhang
tion sensu Holttum & Grimes (1980). Not mono- (2015). Monophyletic (Zhang & Zhang, 2015). Two
phyletic as circumscribed (Almeida et al., 2016). 11 genera and an estimated 22 species.
species.
Hypodematium Kunze, Flora 16(2): 690. 1833. Type:
Sphaerostephanos J.Sm., Gen. Fil. (Hooker): pl.24. Hypodematium onustum Kunze. Circumscription
1840. Type: Sphaerostephanos asplenioides J.Sm. sensu Kramer et al. (1990). Monophyletic (Zhang &
Circumscription sensu Holttum (1971). Not mono- Zhang, 2015). About 20 species.
phyletic as circumscribed (Almeida et al., 2016).
About 185 species. Leucostegia C.Presl, Tent. Pterid.: 94, pl.4, f.11.
1836. Type: Leucostegia immersa C.Presl. Circum-
Stegnogramma Blume, Enum. Pl. Javae 2: 172. scription sensu Kramer (1990c). Monophyletic
1828. Type: Stegnogramma aspidioides Blume. (Zhang & Zhang, 2015). Two species.
Circumscription sensu Almeida et al. (2016).
Includes: Dictyocline T.Moore; Leptogramma J. 45. Family Dryopteridaceae Herter, Revista Sudamer. Bot.
Sm. Monophyletic (Smith & Cranfill, 2002; 9(1): 15. 1949. Circumscription sensu Liu et al. (2016).
He & Zhang, 2012; Almeida et al., 2016). 18 Monophyletic (Liu et al., 2016). Three subfamilies, 26
species. genera, and an estimated 2115 species.
Steiropteris (C.Chr.) Pic.Serm., Webbia 28: 449. Subfamily Polybotryoideae H.M.Liu & X.C.Zhang, Pl.
1973. Lectotype (designated by C.Chr., Kongel. Syst. Evol. 302: 330. 2016. Circumscription sensu Liu
Danske Vidensk. Selsk. Skr., Naturvidensk. Math. et al. (2016). Monophyletic (Moran & Labiak, 2015; Liu
Afd., ser. 7, 1913: 164): Steiropteris deltoidea (Sw.) et al., 2016). Seven genera and an estimated 98
Pic.Serm. ( Polypodium deltoideum Sw.). Cir- species.
cumscription sensu Almeida et al. (2016). In-
cludes: Glaphyropteris Fe e. Monophyletic (Smith Cyclodium C.Presl, Tent. Pterid.: 85, pl.2, f.20–21.
& Cranfill, 2002; He & Zhang, 2012; Almeida et al., 1836. Lectotype (designated by C.Chr., Index Filic.
2016). 22 species. xxv. 1906): Cyclodium meniscioides (Willd.) C.Presl
( Aspidium meniscioides Willd.). Circumscription
Thelypteris Schmidel, Icon. Pl. 3: 45–48, pl.11, 13. sensu Smith (1986) and Moran & Labiak (2015).
1763. Type: Thelypteris palustris Schott ( Monophyletic (Moran & Labiak, 2015). About 10
Acrostichum thelypteris L.). Circumscription species.
sensu Almeida et al. (2016). Monophyletic (Smith
& Cranfill, 2002; He & Zhang, 2012; Almeida et al., Maxonia C.Chr., Smithsonian Misc. Collect. 66(9):
2016). Two species. 3. 1916. Type: Maxonia apiifolia (Sw.) C.Chr. (
Dicksonia apiifolia Sw.). Circumscription sensu
Trigonospora Holttum, Blumea 19(1): 29. 1971. Christensen (1916) and Moran & Labiak (2015).
Type: Trigonospora ciliata (Benth.) Holttum ( Monotypic (Prado & Moran, 2016).
Aspidium ciliatum Benth.). Circumscription sensu
Holttum & Grimes (1980). Monophyly uncertain Olfersia Raddi, Opusc. Sci. 3: 283, t.11, f.b. 1819.
(only one species sampled; Almeida et al., 2016). Type: Olfersia corcovadensis Kaulf. ex Raddi.
Eight species. Circumscription sensu Moran (1986) and Moran

PPG I 587
& Labiak (2015). Monophyletic (Moran & Labiak, Lastreopsis Ching, Bull. Fan Mem. Inst. Biol. Bot. 8
2015). Three species. (4): 157. 1938. Type: Lastreopsis recedens (J.Sm. ex
T.Moore) Ching ( Lastrea recedens J.Sm. ex T.
Polybotrya Humb. & Bonpl. ex Willd., Sp. Pl. 5(1): Moore). Circumscription sensu Tindale (1965) and
99. 1810. Type: Polybotrya osmundacea Humb. & Labiak et al. (2014, 2015b). Includes: Coveniella
Bonpl. ex Willd. Circumscription sensu Moran Tindale. See also Labiak et al. (2015b). Monophy-
(1987) and Moran & Labiak (2015). Monophyletic letic (Labiak et al., 2014). 16 species.
(Moran & Labiak, 2015). 35 species.
Lomagramma J.Sm., J. Bot. (Hooker) 3: 402. 1841.
Polystichopsis (J.Sm.) C.Chr., Verdoorn, Man. Type: Lomagramma pteroides J.Sm. Circumscrip-
Pterid.: 543. 1938. Lectotype (designated by C. tion sensu Moran et al. (2010a). Includes: Cheilo-
Chr., Index Filic. Suppl. 3, 7. 1934): Polystichopsis lepton Fee. Monophyletic (Moran et al., 2010a). 15
pubescens (L.) C.V.Morton ( Polypodium pubes- species.
cens L.). Circumscription sensu Moran & Labiak
(2015) and Prado & Moran (2016). Monophyletic Megalastrum Holttum, Gard. Bull. Singapore 39(2):
(Moran & Labiak, 2015). Eight species. 161. 1986. Type: Megalastrum villosum (L.) Holttum
( Polypodium villosum L.). Circumscription sensu
Stigmatopteris C.Chr., Bot. Tidsskr. 29: 292. 1909. Holttum (1986) and Liu et al. (2016). Monophyletic
Type: Stigmatopteris rotundata (Willd.) C.Chr. ( (Labiak et al., 2014). 91 species.
Aspidium rotundatum Willd.). Circumscription
sensu Moran (1991) and Moran & Labiak (2015, Mickelia R.C.Moran, Labiak & Sundue, Brittonia 62
2016). Monophyletic (Moran & Labiak, 2016). 40 (4): 338. 2010. Type: Mickelia nicotianifolia (Sw.)
species. R.C.Moran, Labiak & Sundue ( Acrostichum
nicotianifolium Sw.). Circumscription sensu Moran
Trichoneuron Ching, Acta Phytotax. Sin. 10(2): 118, et al. (2010a, 2010b). Monophyletic (Moran et al.,
pl. 22. 1965. Type: Trichoneuron microlepioides 2010b). 10 species.
Ching. Circumscription sensu Liu et al. (2016).
Monotypic (Liu et al., 2016). Parapolystichum (Keyserl.) Ching, Sunyatsenia 5:
239. 1940. Type: Parapolystichum effusum (Sw.)
Subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, Ching ( Polypodium effusum Sw.). Circumscrip-
Jermy & Mickel, Fern Gaz. 11: 154. 1975. Circumscription tion sensu Labiak et al. (2014, 2015b). Mono-
sensu Zhang et al. (2013b) and Liu et al. (2016). phyletic (Labiak et al., 2014). See also Labiak
Monophyletic (Liu et al., 2016). 11 genera and an et al. (2015b) and Sundue & Testo (2016). 28
estimated 883 species. species.
Arthrobotrya J.Sm., Hist. Fil.: 141. 1875. Type: Pleocnemia C.Presl, Tent. Pterid.: 183, pl.7,
Arthrobotrya articulata (Fee) J.Sm. ( Polybotrya f.12. 1836. Type: Pleocnemia leuzeana (Gaudich.)
articulata F
ee). Circumscription sensu Moran et al. C.Presl ( Polypodium leuzeanum Gaudich.).
(2010a). Monophyletic (Labiak et al., 2014). Two Circumscription sensu Holttum (1974b) and Liu
species. et al. (2016). Monophyletic (Labiak et al., 2014).
20 species.
Bolbitis Schott, Gen. Fil.: pl.14. 1834. Lectotype
(designated by C.Chr., Index Filic. xxvi. 1906): Rumohra Raddi, Opusc. Sci. 3: 290. 1819. Type:
Bolbitis serratifolia (Mert. ex Kaulf.) Schott ( Rumohra aspidioides Raddi ( Polypodium adian-
Acrostichum serratifolium Mert. ex Kaulf.). Circum- tiforme G.Forst.). Circumscription sensu Rakoton-
scription sensu Moran et al. (2010a). Includes: drainibe (2010), Sundue et al. (2013), and Labiak
Campium C.Presl; Cyrtogonium J.Sm.; Edanyoa et al. (2014). Monophyletic (Labiak et al., 2014).
Copel.; Egenolfia Schott; Heteroneuron Fe e; Eight species.
Jenkinsia Hook.; Poecilopteris C.Presl. Monophy-
letic (Moran et al., 2010a, 2010b). About 80 Teratophyllum Mett. ex Kuhn, Ann Mus. Bot.
species. Lugduno-Batavi 4(10): 296. 1869. Lectotype
(designated by Holttum, Gard. Bull. Straits Settlem.
Elaphoglossum Schott ex J.Sm., J. Bot. (Hooker) 4: 5: 277. 1932): Teratophyllum aculeatum (Blume)
148. 1842. Lectotype (designated by J.Sm., Hist. Fil.: Mett. ex Kuhn ( Lomaria aculeata Blume).
125. 1875): Elaphoglossum conforme (Sw.) J.Sm. ( Circumscription sensu Moran et al. (2010a). Mono-
Acrostichum conforme Sw.). Circumscription sensu phyletic (Moran et al., 2010a). 13 species.
Moran et al. (2010a). Includes: Aconiopteris C.Presl;
Dictyoglossum Sm.; Hymenodium Fe e; Microsta- Subfamily Dryopteridoideae Link, Fil. Spec.: 116. 1841
phyla C.Presl; Peltapteris Link; Rhipidopteris Schott (Dryopterideae). Circumscription sensu Liu et al. (2016).
ex Fee. Monophyletic (Rouhan et al., 2004; Lo riga Monophyletic (Liu et al., 2016). Six genera and an
et al., 2014). About 600 species. estimated 1128 species.

Arachniodes Blume, Enum. Pl. Javae 2: 241. 1828. Ptilopteris Hance; Sorolepidium Christ. Mono-
Type: Arachniodes aspidioides Blume. Circumscrip- phyletic (Le Pechon et al., 2016). About 500
tion sensu He et al. (2013) and Liu et al. (2016). species.
Includes: Byrsopteris C.V.Morton; Leptorumohra
(H.Ito) H.Ito; Lithostegia Ching; Phanerophlebiop- Subfamily placement uncertain.
sis Ching. Assumed to be monophyletic (Liu et al.,
2016). 60 species. Aenigmopteris Holttum, Blumea 30: 3. 1984. Type:
Aenigmopteris dubia (Copel.) Holttum ( Dryop-
Ctenitis (C.Chr.) C.Chr., Man. Pteridol.: 544. 1938. teris dubia Copel.). Circumscription sensu Kramer
Lectotype (designated by Ching, Bull. Fan et al. (1990). No molecular data available. Five
Mem. Inst. Biol. Bot. 8(5): 275. 1938): Ctenitis species.
distans (Brack.) Ching ( Lastrea distans
Brack.). Circumscription sensu Wang et al. Dryopolystichum Copel., Ann. Cryptog. Phytopa-
(2014), Liu et al. (2016), and Duan et al. (2017). thol. 5 [Gen. Fil.]: 125. 1947. Type: Dryopolysti-
Includes: Ataxipteris Holttum; Pseudotectaria chum phaeostigma (Ces.) Copel. ( Aspidium
Tardieu. Monophyletic (Wang et al., 2014; phaeostigma Ces.). Circumscription sensu Chris-
Hennequin et al., pers. comm.). About 125 tenhusz et al. (2011). Monotypic (Christenhusz
species. et al., 2011).
Cyrtomium C.Presl, Tent. Pterid.: 86, pl.2, f.26. 46. Family Nephrolepidaceae Pic.Serm., Webbia 29(1):
1836. Lectotype (designated by J.Sm., Hist. Fil.: 8–11. 1975. Circumscription sensu Hovenkamp &
204. 1875): Cyrtomium falcatum (L.f.) C.Presl ( Miyamoto (2012). Family consists of a single genus
Polypodium falcatum L.f.). Circumscription sensu and an estimated 19 species.
Lu et al. (2007), Zhang et al. (2013b), and Liu et al.
(2016). Assumed to be monophyletic (Lu et al., Nephrolepis Schott, Gen. Fil.: 1, pl.3. 1834.
2007). 35 species. Lectotype (designated by J.Sm., Hist. Fil. 226.
1875): Nephrolepis exaltata (L.) Schott ( Poly-
Dryopteris Adans., Fam. Pl. 2: 20. 1763. Type: podium exaltatum L.). Circumscription sensu
Dryopteris filix-mas (L.) Schott ( Polypodium Hennequin et al. (2010). Monophyletic (Henne-
filix-mas L.). Circumscription sensu Zhang & quin et al., 2010; Zhang et al., 2016). 19 species.
Zhang (2012), Zhang et al. (2012), and Zhang
et al. (2013b). Includes: Acrophorus C.Presl; 47. Family Lomariopsidaceae Alston, Taxon 5(2): 25. 1956.
Acrorumohra (H.Ito) H.Ito; Adenoderris J.Sm, in Circumscription sensu Christenhusz et al. (2013).
part; Arthrobotrys (C.Presl) Lindl.; Diacalpe Monophyletic (Christenhusz et al., 2013; Zhang et al.,
Blume; Dichasium (A.Braun) F ee; Diclisodon T. 2016). Four genera and an estimated 69 species.
Moore; Dryopsis Holttum & P.J.Edwards; Filix
Seg.; Lophodium Newman; Nephrodium Marthe Cyclopeltis J.Sm., Bot. Mag. 72: 36. 1846. Type:
ex Michx.; Nothoperanema (Tagawa) Ching; Cyclopeltis semicordata (Sw.) J.Sm. ( Polypo-
Peranema D.Don; Pycnopteris T.Moore; Revwatt- dium semicordatum Sw.). Circumscription sensu
sia D.L.Jones; Stenolepia Alderw. Monophyletic Kramer et al. (1990). Monophyletic (Zhang &
(Zhang et al., 2012; Sessa et al., 2012; Zhang & Zhang, 2015). Six species.
Zhang, 2012; McKeown et al., 2012; Kuo et al.,
2016a). About 400 species. Dracoglossum Christenh., Thaiszia 17(1–2): 3–5.
2007. Type: Dracoglossum plantagineum (Jacq.)
Phanerophlebia C.Presl, Tent. Pterid.: 84, pl.2, Christenh. ( Polypodium plantagineum Jacq.).
f.19. 1836. Type: Phanerophlebia nobilis (Schltdl. & Circumscription sensu Christenhusz (2007).
Cham.) C.Presl ( Aspidium nobile Schltdl. & Monophyletic (Zhang et al., 2016). Two species.
Cham.). Circumscription sensu Yatskievych et al.
(1988) and Liu et al. (2016). Includes: Amblia C. Lomariopsis Fe e, Mem. Foug., 2. Hist. Acrostich.:
Presl. Monophyletic (Yatskievych et al., 1988). 10. 1845. Lectotype (designated by J.Sm., Hist.
Eight species. e (
Fil.: 140. 1875): Lomariopsis sorbifolia (L.) Fe
Acrostichum sorbifolium L.). Circumscription
Polystichum Roth, Tent. Fl. Germ. 3(1): 31, 69–70. sensu Rouhan et al. (2007) and Moran (2000).
1800. Lectotype (designated by J.Sm., Hist. Fil.: Monophyletic (Rouhan et al., 2007; Li et al.,
217. 1875): Polystichum aculeatum (L.) Roth 2009). About 60 species.
( Polypodium aculeatum L.). Circumscription
sensu Zhang et al. (2013b) and Le Pe chon et al. Thysanosoria A.Gepp, Fl. Arfak Mts. (Gibbs): 193–
(2016). Includes: Acropelta T.Nakai; Adenoderris 194, pl.4. 1917. Type: Thysanosoria dimorphophylla
J.Sm, in part; Cyrtogonellum Ching; Cyrtomidic- A.Gepp. Circumscription sensu Kramer (1990h).
tyum Ching; Hemesteum H.Le v.; Hypopeltis Genus has yet to be included in a phylogenetic
Michx.; Papuapteris C.Chr.; Plecosorus Fe e; analysis. Monotypic.

PPG I 589
48. Family Tectariaceae Panigrahi, J. Orissa Bot. Soc. 8: Triplophyllum Holttum, Kew Bull. 41(2): 239. 1986.
41. 1986. Circumscription sensu Zhang et al. (2016). Type: Triplophyllym protensum (Afzel. ex Sw.)
Monophyletic (Zhang et al., 2016). Seven genera and Holttum ( Aspidium protensum Afzel. ex Sw.).
an estimated 250 species. Circumscription sensu Holttum (1986). Monophy-
letic (Prado & Moran, 2008). About 20 species.
Arthropteris J.Sm. ex Hook.f., Fl. Nov.-Zel. 2: 43.
1855. Type: Arthropteris tenella (G.Forst.) J.Sm. ex 49. Family Oleandraceae Ching ex Pic.Serm., Webbia 20
Hook.f. ( Polypodium tenellum G.Forst.). Cir- (2): 745. 1965. Circumscription sensu Smith et al.
cumscription sensu Liu et al. (2013). Includes: (2006b). Family consists of a single genus and an
Psammiosorus C.Chr. Monophyletic (Liu et al., estimated 15 species.
2013; Zhang et al., 2016). About 15 species.
Oleandra Cav., Anales Hist. Nat. 1(2): 115. 1799.
Draconopteris Li Bing Zhang & Liang Zhang, Type: Oleandra neriiformis Cav. Circumscription
Taxon 65(4): 732, f.1a–b, f.3a–d. 2016. Type: sensu Kramer (1990n). Monophyletic (Zhang &
Draconopteris draconoptera (D.C.Eaton) Li Bing Zhang, 2015). At least 15 species.
Zhang & Liang Zhang ( Aspidium draconopterum
D.C.Eaton). Circumscription sensu Zhang et al. 50. Family Davalliaceae M.R.Schomb., Reis. Br.-Guiana 3:
(2016). Monotypic. 883. 1848. Circumscription sensu Smith et al.
(2006b). Family consists of a single genus and an
Hypoderris R.Br. in Wall., Pl. Asiat. Rar. (Wallich) 1: estimated 65 species.
16. 1829. Lectotype (designated by J.Sm. in Hook.
Gen. Fil.: t.1. 1838.): Hypoderris brownii J.Sm. Davallia Sm., Me m. Acad. Roy. Sci. (Turin) 5: 414,
Circumscription sensu Moran et al. (2014). pl. 9, f. 6. 1793. Lectotype (designated by Fe e,
Monophyletic (Moran et al., 2014; Zhang et al., Mem. Foug., 5. Gen. Filic.: 328. 1852): Davallia
2016). Three species. canariensis (L.) Sm. ( Trichomanes canariense L.).
Circumscription sensu Tsutsumi et al. (2016).
Malaifilix Li Bing Zhang & Schuettp., Taxon 65(4): Includes: Araiostegia Copel.; Araiostegiella M.Kato
733, f.1e–f, f.3e–f. 2016. Type: Malaifilix grandi- & Tsutsumi; Davallodes (Copel.) Copel.; Humata
dentata (Ces.) Li Bing Zhang & Schuettp. ( Cav.; Scyphularia Fe e; Wibelia Bernh. Monophy-
Polypodium dilatatum var. grandidentatum Ces.). letic (Tsutsumi & Kato, 2006; Tsutsumi et al.,
Circumscription sensu Zhang et al. (2016). Pre- 2016). About 65 species.
sumably monotypic.
51. Family Polypodiaceae J.Presl & C.Presl, Delic. Prag.:
Pteridrys C.Chr. & Ching, Bull. Fan Mem. Inst. Biol. 159. 1822. Circumscription sensu Smith et al. (2006b).
Bot. 5(3): 129. 1934. Type: Pteridrys syrmatica Includes: Loxogrammaceae Ching ex Pic.Serm.;
(Willd.) C.Chr. & Ching ( Aspidium syrmaticum Grammitidaceae Newman; Platyceriaceae Ching;
Willd.). Circumscription sensu Christensen & Drynariaceae Ching. Monophyletic (Schuettpelz &
Ching (1934). Monophyletic (Zhang et al., 2016). Pryer, 2007). Six subfamilies, 65 genera, and an
About 10 species. estimated 1652 species.
Tectaria Cav., Anales Hist. Nat. 1(2): 115. 1799. Subfamily Loxogrammoideae H.Schneid. in Christen-
Type: Tectaria trifoliata (L.) Cav. ( Polypodium husz et al., Phytotaxa 19: 18. 2011. Circumscrip-
trifoliatum L.). Circumscription sensu Zhang et al. tion equivalent to “loxogrammoid clade“ in Kreier
(2016). Includes: Amphiblestra C.Presl; Aspidium & Schneider (2006a). Monophyletic (Kreier &
Sw.; Bathmium C.Presl ex Link; Camptodium Fe e; Schneider, 2006a). Two genera and an estimated
Chlamydogramme Holttum; Cionidium T.Moore; 32 species.
Ctenitopsis Ching ex Tardieu & C.Chr.; Dictyox-
iphium Hook.; Dryomenis Fe e ex J.Sm.; Fadyenia Dictymia J.Sm., Companion Bot. Mag. 72: 16. 1846.
Hook.; Grammatosorus Regel; Hemigramma Type: Dictymia attenuata J.Sm. Circumscription
Christ; Heterogonium C.Presl; Lenda Koidz.; Luers- sensu Kreier & Schneider (2006a). Monophyletic
senia Kuhn ex Luerssen; Microbrochis C.Presl; (Kreier & Schneider, 2006a). No published generic
Phlebiogonium Fe e; Podopeltis Fe
e; Polydictyum revision available. About two species.
C.Presl; Psomiocarpa C.Presl; Quercifilix Copel.;
Sagenia C.Presl; Stenosemia C.Presl; Tectaridium Loxogramme (Blume) C.Presl, Tent. Pterid.:
Copel. Monophyletic (Zhang et al., 2016), but 214–215, pl.9, f.8. 1836. Lectotype (designated
molecular data is lacking for Amphiblestra, by J.Sm., Hist. Fil.: 156. 1875): Loxogramme
Camptodium, Chlamydogramme, Dryomenis, lanceolata (Sw.) C.Presl ( Grammitis lanceolata
Grammatosorus, Lenda, Luerssenia, Microbrochis, Sw.). Circumscription sensu Kreier & Schneider
Phlebiogonium, and Polydictyum (tentatively in- (2006a). Includes: Anarthropteris Copel. Mono-
cluded here based on morphology). About 200 phyletic (Kreier & Schneider, 2006a). No pub-
species. lished generic revision available. 30 species.

Subfamily Platycerioideae B.K.Nayar, Taxon 19: 233. Selliguea Bory, Dict. Class. Hist. Nat. 6: 587–588.
1970. Circumscription sensu Hennipman et al. (1990). 1824. Type: Selliguea feei Bory. Circumscription
Monophyletic (Kreier & Schneider, 2006b). Two sensu Schneider et al. (2008). Includes: Crypsinus
genera and an estimated 70 species. C.Presl; Himalayopteris W.Shao & S.G.Lu; Phyma-
topsis J.Sm.; Pichisermollodes Fraser-Jenk. &
Platycerium Desv., M
em. Soc. Linn. Paris 6(3): 213. Challis. Not monophyletic as circumscribed
1827. Lectotype (designated by Pfeiff., Nomencl. (Schneider et al., 2008; Li et al., 2012a); Selliguea
Bot. 2: 746. 1873): Platycerium alcicorne Desv. appears to be polyphyletic relative to Arthrome-
Circumscription sensu Hennipman et al. (1990). ris, Gymnogrammitis, and Polypodiopteris, but
Monophyletic (Kreier & Schneider, 2006b). 18 additional study is needed. At least 85 species.
species.
Subfamily Microsoroideae B.K.Nayar, Taxon 19(2):
Pyrrosia Mirb., Hist. Nat. Veg. 3: 471. 1803. Type: 233. 1970. Circumscription equivalent to “microsor-
Pyrrosia chinensis Desv. Circumscription sensu oids” in Kreier et al. (2008b). Monophyletic
Hennipman et al. (1990). Includes: Drymoglossum (Schneider et al., 2004c). 12 genera and an estimated
C.Presl. Assumed to be monophyletic (Kreier & 183 species.
Schneider, 2006b), but paraphyly relative to
Platycerium has not yet been definitively ex- Goniophlebium (Blume) C.Presl, Tent. Pterid.: 185,
cluded. At least 52 species. € dl-
pl.7, f.13–14. 1836. Lectotype (designated by Ro
Linder, Zijlstra & Tryon, Taxon 39(1): 105. 1990):
Subfamily Drynarioideae Crabbe, Jermy & Mickel, Goniophlebium subauriculatum (Blume) C.Presl (
Fern Gaz. 11: 156. 1975. Circumscription sensu Polypodium subauriculatum Blume). Circumscrip-
Christenhusz et al. (2011). Monophyletic (Schneider tion sensu Kreier et al. (2008b). Includes:
et al., 2008). Six genera and an estimated 148 Metapolypodium Ching; Polypodiastrum Ching;
species. Polypodiodes Ching. Monophyletic (Kreier et al.,
2008b). 25 species.
Aglaomorpha Schott, Gen. Fil.: t.19. 1836. Type:
Aglaomorpha meyeniana Schott. Circumscription Lecanopteris Reinw. ex Blume, Enum. Pl. Javae 2:
newly proposed in this classification. Includes: 120. 1828. Type: Lecanopteris carnosa (Reinw.)
Christiopteris Copel., Drynaria J.Sm., Dryosta- Blume ( Onychium carnosum Reinw.). Circum-
chyum J.Sm., Merinthosorus Copel., Photinopteris scription sensu Hennipman et al. (1990). Mono-
J.Sm., Pseudodrynaria C.Chr., Thayeria Copel. phyletic (Kreier et al., 2008b). 13 species.
Monophyletic (Schneider et al., 2008). 50 species.
Lemmaphyllum C.Presl, Abh. Koenigl. B€ ohm. Ges.
Arthromeris (T.Moore) J.Sm., Hist. Fil.: 110. 1875. Wiss. Ser. 5 6: 517–518. 1851. Lectotype (desig-
Type: Arthromeris wallichiana (Spreng.) Ching ( nated by Copel., Ann. Cryptog. Phytopathol. 5
Polypodium wallichianum Spreng.). Circumscrip- [Gen. Fil.]: 189. 1947): Lemmaphyllum spathulatum
tion sensu Hennipman et al. (1990). Assumed to C.Presl. Circumscription sensu Wang et al.
be monophyletic (Schneider et al., 2008), but (2010). Includes: Caobangia A.R.Sm. & X.C.Zhang;
comprehensive analysis is lacking to date. Lepidogrammitis Ching; Weatherbya Copel.
Perhaps a few more species, if a narrower Monophyletic (Wang et al., 2010). At least two
species concept is used. Eight species. species.
Gymnogrammitis Griff., Icon. Pl. Asiat. 2: t.129, f.1. Lepidomicrosorium Ching & K.H.Shing, Bot. Res.
1849. Type: Gymnogrammitis dareiformis (Hook.) Academia Sinica 1(1): 1–14, pl.1–5. 1983. Type:
Ching ex Tardieu & C.Chr. ( Polypodium dare- Lepidomicrosorium subhastatum (Baker) Ching (
iforme Hook.). Circumscription sensu Kramer Polypodium subhastatum Baker). Circumscription
(1990c). Monotypic. sensu Wang et al. (2010). Apparently monotypic
(Wang et al., 2010).
Paraselliguea Hovenkamp, Blumea 45(2): 376.
2000. Type: Paraselliguea leucophora (Baker) Lepisorus (J.Sm.) Ching, Bull. Fan Mem. Inst. Biol.
Hovenkamp ( Polypodium leucophorum Baker). Bot. 4(3): 47, 56–58. 1933. Type: Lepisorus nudus
Circumscription sensu Hovenkamp (1997). Mono- (Hook.) Ching ( Pleopeltis nuda Hook.). Circum-
typic. scription sensu Wang et al. (2010). Includes:
Belvisia Mirb.; Drymotaenium Makino. Monophy-
Polypodiopteris C.F.Reed, Amer. Fern J. 38: 87. letic (Wang et al., 2010). 80 species.
1948. Type: Polypodiopteris proavita (Copel.) C.F.
Reed ( Polypodium proavitum Copel.). Circum- Leptochilus Kaulf., Enum. Filic.: 147, pl.1, f.10. 1824.
scription sensu Hennipman et al. (1990). As- Type: Leptochilus axillaris (Cav.) Kaulf. ( Acros-
sumed to be monophyletic, but only one species tichum axillare Cav.). Circumscription sensu Kreier
has been sampled to date. Three species. et al. (2008b). Includes: Colysis C.Presl; Kontumia

PPG I 591
S.K.Wu & K.L.Phan. Monophyletic (Kreier et al., Le

on (1995), Schneider et al. (2004c), and Kreier
2008b). 10 species. et al. (2007). Includes: Hyalotricha Copel.;
Hyalotrichopteris W.H.Wagner. Monophyletic
Microsorum Link, Hort. Berol. 2: 110. 1833. Type: (Schneider et al., 2004c). About 50 species.
Microsorum irregulare Link. Circumscription sensu
Kreier et al. (2008b). Includes: Dendroconche Microgramma C.Presl, Tent. Pterid.: 213. 1836.
Copel.; Kaulinia B.K.Nayar; Phymatosorus Pic. Type: Microgramma persicariifolia (Schrad.) C.
Serm. Not monophyletic as circumscribed (Kreier Presl ( Polypodium persicariifolium Schrad.).
et al., 2008b); Microsorum is apparently para- Circumscription sensu Salino et al. (2008).
phyletic relative to Lecanopteris, Leptochilus, Includes: Solanopteris Copel. Monophyletic (Sal-
Neocheiropteris, Neolepisorus, and Tricholepi- ino et al., 2008). A generic revision is lacking.
dium, but further study is needed. 40 species. About 30 species.
Neocheiropteris Christ, Bull. Soc. Bot. France 52 Niphidium J.Sm., Hist. Fil.: 99. 1875. Type:
(Mem. 1): 21. 1905. Type: Neocheiropteris palma- Niphidium americanum (Hook.) J.Sm. ( Polypo-
topedata (Baker) Christ ( Polypodium palmato- dium americanum Hook.). Circumscription sensu
pedatum Baker). Circumscription sensu Wang Hennipman et al. (1990). Monophyletic (Salino
et al. (2010). Monophyletic (Kreier et al., 2008b; et al., 2008). 10 species.
Wang et al., 2010; Du & Cheng, 2011). Two species.
Pecluma M.G.Price, Amer. Fern J. 73(4): 109. 1983.
Neolepisorus Ching, Bull. Fan Mem. Inst. Biol. Bot. Type: Pecluma pectinata (L.) M.G.Price ( Poly-
10(1): 11–12. 1940. Type: Neolepisorus ensatus podium pectinatum L.). Circumscription sensu
(Thunb.) Ching ( Polypodium ensatum Thunb.). Assis et al. (2016). Monophyletic (Assis et al.,
Circumscription sensu Wang et al. (2010). Mono- 2016). 40 species.
phyletic (Wang et al., 2010). The generic affinities
have not been evaluated for all species possibly Phlebodium (R.Br.) J.Sm., J. Bot. (Hooker) 4: 58.
belonging here. At least five species. 1841. Lectotype (designated by J.Sm., Hist. Fil.:
94. 1875): Phlebodium aureum (L.) J.Sm. (
Paragramma (Blume) T.Moore, Index Filic. xxxii. Polypodium aureum L.). Circumscription sensu
1857. Lectotype (designated by J.Sm., Hist. Fil.: Mickel & Smith (2004). Monophyletic (Assis et al.,
114. 1875): Paragramma longifolia (Blume) T. 2016). Four species.
Moore ( Grammitis longifolia Blume). Circum-
scription sensu Wang et al. (2010). Only the type Pleopeltis Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4,
species has been sampled to date; the second 5: 211. 1810. Type: Pleopeltis angusta Humb. &
species is highly distinct and may ultimately be Bonpl. ex Willd. Circumscription sensu Smith &
resolved elsewhere. Perhaps two species. Tejero-Dıez (2014). Includes: Cheilogramma
Maxon; Dicranoglossum J.Sm.; Eschatogramme
Thylacopteris Kunze ex J.Sm., Hist. Fil.: 87. 1875. Trevis.; Lepicystis (J.Sm.) J.Sm.; Marginaria Bory;
Type: Thylacopteris papillosa (Blume) J.Sm. ( Marginariopsis C.Chr.; Microphlebodium L.D.
Polypodium papillosum Blume). Circumscription Gomez; Neurodium F ee; Paltonium C.Presl; Pseu-
sensu Hennipman et al. (1990). Assumed to be docolysis L.D.Gomez. Monophyletic (Otto et al.,
monophyletic, but only the type species has been 2009). In need of a generic revision. About 90
sampled to date. Two species. species.
Tricholepidium Ching, Acta Phytotax. Geobot. 29 Pleurosoriopsis Fomin, Izv. Kievsk. Bot. Sada 11: 8.
(1–5): 41. 1978. Type: Tricholepidium normale (D. 1930. Type: Pleurosoriopsis makinoi (Maxim. ex
Don) Ching ( Polypodium normale D.Don). Makino) Fomin ( Gymnogramma makinoi
Circumscription sensu Wang et al. (2010). Mono- Maxim. ex Makino). Circumscription sensu Hen-
typic (Wang et al., 2010), but in need of further nipman et al. (1990). Monotypic (Assis et al.,
evaluation. 2016).
Subfamily Polypodioideae Sweet, Hort. Brit.: 460. Polypodium L., Sp. Pl. 2: 1082. 1753. Lectotype
1826. Newly circumscribed in this classification. (designated by J.Sm., Hist. Fil.: 88. 1875): Poly-
Paraphyletic relative to the Grammitidoideae podium vulgare L. Circumscription sensu Sigel
(Schneider et al., 2004c). Nine genera and an et al. (2014) and Assis et al. (2016). Presumably
estimated 305 species. monophyletic (Sigel et al., 2014; Assis et al., 2016).
Perhaps 40 species.
Campyloneurum C.Presl, Tent. Pterid.: 189. 1836.
Lectotype (designated by J.Sm., Hist. Fil.: 95. Serpocaulon A.R.Sm., Taxon 55(4): 924–927, f. 3-
1875): Campyloneurum repens (Aubl.) C.Presl ( 4. 2006. Type: Serpocaulon loriceum (L.) A.R.Sm.
Polypodium repens Aubl.). Circumscription sensu ( Polypodium loriceum L.). Circumscription

sensu Smith et al. (2006a). Monophyletic (Smith Chrysogrammitis Parris, Kew Bull. 3(4): 909.
et al., 2006a; Kreier et al., 2008a). About 40 1998. Type: Chrysogrammitis glandulosa (J.Sm.)
species. Parris ( Ctenopteris glandulosa J.Sm.). Circum-
scription sensu Parris (1998) and Sundue et al.
Subfamily Grammitidoideae Parris & Sundue, this (2014). Monophyletic (Sundue et al., 2014). Two
classification (see below). Circumscription equivalent species.
to “grammitids” in Sundue et al. (2014). Monophy-
letic (Ranker et al., 2004; Schneider et al., 2004c; Cochlidium Kaulf., Berlin. Jahrb. Pharm. Verbun-
Sundue et al., 2014). 33 genera and an estimated 911 denen Wiss. 21: 36. 1820. Lectotype (designated
species. by Copel., Ann. Cryptog. Phytopathol. 5 [Gen.
Fil.]: 213. 1947): Cochlidium graminoides (Sw.)
Acrosorus Copel., Philipp. J. Sci. 1(Suppl. 2): 158– Kaulf. ( Acrostichum graminoides Sw.). Circum-
159. 1906. Lectotype (designated by Copel., Univ. scription sensu Bishop (1978) and Sundue et al.
Calif. Publ. Bot. 16: 108. 1929): Acrosorus exaltatus (2014). Includes: Xiphopteris Kaulf.; Micropteris
(Copel.) Copel. ( Davallia exaltata Copel.). Desv.; Pleurogramme Blume. Monophyletic, but
Circumscription sensu Parris (1990) and Sundue nested within Grammitis (Sundue et al., 2014).
et al. (2014). Assumed to be monophyletic, but Grammitis s.s. was found to be paraphyletic with
only one species sampled to date (Sundue et al., regards to Cochlidium in Sundue et al. (2014), but
2014). Nine species. the type, Grammitis marginella, has not been
sampled. We prefer to maintain both genera as
Adenophorus Gaudich., Ann. Sci. Nat. (Paris) 3: they are circumscribed until additional data
508. 1824. Lectotype (designated by J.Sm., Hist. becomes available. 18 species.
Fil.: 187. 1875): Adenophorus tripinnatifidus Gau-
dich. Circumscription sensu Ranker (2008) and Ctenopterella Parris, Gard. Bull. Singapore 58(2):
Sundue et al. (2014). Includes: Amphoradenium 234. 2007. Type: Ctenopterella blechnoides (Grev.)
Desv. Monophyletic (Sundue et al., 2014). 10 Parris ( Grammitis blechnoides Grev.). Circum-
species. scription sensu Parris (2007) and Sundue et al.
(2014). Only one species sampled to date
Alansmia M.Kessler, Moguel, Sundue & Labiak, (Sundue et al., 2014). 24 species.
Brittonia 63(2): 238. 2011. Type: Alansmia lanigera
(Desv.) Moguel & M.Kessler ( Polypodium Dasygrammitis Parris, Gard. Bull. Singapore 58(2):
lanigerum Desv.). Circumscription sensu Kessler 238. 2007. Type: Dasygrammitis mollicoma (Nees
et al. (2011) and Sundue et al. (2014). Monophy- & Blume) Parris ( Polypodium mollicomum Nees
letic (Sundue et al., 2014). 26 species. & Blume). Circumscription sensu Parris (2007)
and Sundue et al. (2014). Monophyletic (Sundue
Archigrammitis Parris, Fern Gaz. 19(4): 135–136. et al., 2014). 12 species.
2013. Type: Archigrammitis friderici-et-pauli
(Christ) Parris ( Polypodium friderici-et-pauli Enterosora Baker, Timehri 5: 218. 1886. Type:
Christ). Circumscription sensu Parris (2013). No Enterosora campbellii Baker. Circumscription
molecular data available. Seven species. sensu Bishop & Smith (1992) pro parte and
Sundue et al. (2014). Monophyletic, but nested
Ascogrammitis Sundue, Brittonia 62(4): 361. 2010. within Zygophlebia (Sundue et al., 2014). This
Type: Ascogrammitis athyrioides (Hook.) Sundue could be resolved by placing Zygophlebia in
( Polypodium athyrioides Hook.). Circumscrip- synonymy. However, this would be premature
tion sensu Sundue (2010) and Sundue et al. since the type of Enterosora has not yet been
(2014). Monophyletic (Sundue et al., 2014). 17 sampled and its placement is not certain. 11
species. species.
Calymmodon C.Presl, Tent. Pterid.: 203–204, pl.9, Galactodenia Sundue & Labiak, Syst. Bot. 37(2):
f.1. 1836. Type: Calymmodon cucullatus (Nees & 340. 2012. Type: Galactodenia delicatula (M.
Blume) C.Presl ( Polypodium cucullatum Nees & Martens & Galeotti) Sundue & Labiak ( Poly-
Blume). Circumscription sensu Parris (1990) and podium delicatulum M.Martens & Galeotti).
e.
Sundue et al. (2014). Includes: Plectopteris Fe Circumscription sensu Sundue et al. (2012,
Monophyletic (Sundue et al., 2014). 65 species. 2014). Monophyletic (Sundue et al., 2014). Five
species.
Ceradenia L.E.Bishop, Amer. Fern J. 78(1): 2. 1988.
Type: Ceradenia curvata (Sw.) L.E.Bishop ( Grammitis Sw., J. Bot. (Schrader) 2: 3, 17. 1801.
Polypodium curvatum Sw.). Circumscription Lectotype (designated by C.Chr., Index Filic. xlix.
sensu Bishop (1988) and Sundue et al. (2014). 1906): Grammitis marginella (Sw.) Sw. ( Poly-
Monophyletic (Sundue et al., 2014). About 73 podium marginellum Sw.). Circumscription sensu
species. Bishop (1977) pro parte and Sundue et al. (2014).

PPG I 593
Grammitis s.s. is paraphyletic relative to Cochli- Notogrammitis Parris, New Zealand J. Bot. 50(4):
dium (Sundue et al., 2014), but the type, G. 465. 2012. Type: Notogrammitis billardierei
marginella, has not been sampled. We prefer to (Willd.) Parris ( Grammitis billardierei Willd.).
maintain both genera as they are circumscribed Circumscription sensu Perrie and Parris (2012).
until additional data become available. 43 spe- Monophyletic (Sundue et al., 2014). 16 species.
cies.
Oreogrammitis Copel., Philipp. J. Sci. 12: 64. 1917.
Lellingeria A.R.Sm. & R.C.Moran, Amer. Fern J. 81 Type: Oreogrammitis clemensiae Copel. Circum-
(3): 76. 1991. Type: Lellingeria apiculata (Kunze ex scription sensu Parris (2007) and Sundue et al.
Klotzsch) A.R.Sm. & R.C.Moran ( Polypodium (2014). Not monophyletic and in need of
apiculatum Kunze ex Klotzsch). Circumscrip- redefinition (Sundue et al., 2014). Themelium is
tion sensu Labiak (2013) and Sundue et al. monophyletic in Sundue et al. (2014), but nested
(2014). Monophyletic (Sundue et al., 2014). 49 within a clade containing polyphyletic Oreogram-
species. mitis and Radiogrammitis. The types of Oreog-
rammitis and Themelium remain to be sampled,
Leucotrichum Labiak, Taxon 59(3): 915. 2010. and thus we recommend maintaining these
Type: Leucotrichum organense (Gardner) Labiak genera pending further research. 156 species.
( Grammitis organensis Gardner). Circumscrip-
tion sensu Labiak et al. (2010), Rouhan et al. Prosaptia C.Presl, Tent. Pterid.: 165–166, pl.6, f.19,
(2012), and Sundue et al. (2014). Monophyletic 25. 1836. Lectotype (designated by J.Sm., Hist.
(Sundue et al., 2014). Six species. Fil.: 86. 1875): Prosaptia contigua (G.Forst.) C.
Presl ( Trichomanes contiguum G.Forst.). Cir-
Lomaphlebia J.Sm., Hist. Fil.: 182. 1875. Type: cumscription sensu Parris (2010a, 2010b) and
Lomaphlebia linearis (Sw.) J.Sm. ( Grammitis Sundue et al. (2014). Includes: Ctenopteris Blume
linearis Sw.). Circumscription sensu Sundue et al. ex Kunze; Cryptosorus Fe e. Monophyletic (Sun-
(2014). Assumed to be monophyletic, but only due et al., 2014). 87 species.
one species sampled to date (Sundue et al.,
2014). Two species. Radiogrammitis Parris, Gard. Bull. Singapore 58
(2): 240. 2007. Type: Radiogrammitis setigera
Luisma M.T.Murillo & A.R.Sm., Novon 13(3): 313– (Blume) Parris ( Polypodium setigerum Blume).
316, f. 1. 2003. Type: Luisma bivascularis M.T. Circumscription sensu Parris (2007) and Sundue
Murillo & A.R.Sm. Circumscription sensu Murillo et al. (2014). Not monophyletic and in need of
& Smith (2003). Monotypic. redefinition (Sundue et al., 2014). Themelium
is monophyletic in Sundue et al. (2014), but
Melpomene A.R.Sm. & R.C.Moran, Novon 2(4): nested within a clade containing polyphyletic
426. 1992. Type: Melpomene moniliformis (Lag. ex Oreogrammitis and Radiogrammitis. The types of
Sw.) A.R.Sm. & R.C.Moran ( Polypodium mon- Oreogrammitis and Themelium remain to be
iliforme Lag. ex Sw.). Circumscription sensu sampled, and thus we recommend maintain-
Lehnert (2013) and Sundue et al. (2014). Mono- ing these genera pending further research. 36
phyletic (Sundue et al., 2014). 29 species. species.
Micropolypodium Hayata, Bot. Mag. (Tokyo) 42 Scleroglossum Alderw., Bull. Jard. Bot. Buiten-
(449): 341. 1928. Type: Micropolypodium pseudo- zorg, ser. 2, 2(7): 37–39. 1912. Lectotype (desig-
trichomanoides (Hayata) Hayata ( Polypodium nated by Copel., Ann. Cryptog. Phytopathol. 5
pseudotrichomanoides Hayata). Circumscription [Gen. Fil.]: 213. 1947): Scleroglossum pusillum
sensu Smith (1992) pro parte, Hirai et al. (2011), (Blume) Alderw. ( Vittaria pusilla Blume).
and Sundue et al. (2014). Monophyletic (Sundue Circumscription sensu Parris (1990) and Sundue
et al., 2014). Three species. et al. (2014). Includes: Nematopteris Alderw.
Monophyletic (Sundue et al., 2014). Seven
Moranopteris R.Y.Hirai & J.Prado, Taxon 60(4): species.
1127. 2011. Type: Moranopteris basiattenuata (Jen-
man) R.Y.Hirai & J.Prado ( Polypodium basiatte- Stenogrammitis Labiak, Brittonia 63(1): 141, f.1A–
nuatum Jenman). Circumscription sensu Hirai M, 2A–F. 2011. Type: Stenogrammitis myosuroides
et al. (2011) and Sundue et al. (2014). Monophy- (Sw.) Labiak ( Polypodium myosuroides Sw.).
letic (Sundue et al., 2014). 31 species. Circumscription sensu Labiak (2011) and Sundue
et al. (2014). Monophyletic (Sundue et al., 2014).
Mycopteris Sundue, Brittonia 66(2): 175, f.1–2. 31 species.
2013. Type: Mycopteris taxifolia (L.) Sundue (
Polypodium taxifolium L.). Circumscription sensu Terpsichore A.R.Sm., Novon 3(4): 479. 1993. Type:
Sundue (2013) and Sundue et al. (2014). Mono- Terpsichore asplenifolia (L.) A.R.Sm. ( Polypo-
phyletic (Sundue et al., 2014). 21 species. dium asplenifolium L.). Circumscription sensu

Smith (1993) pro parte and Sundue et al. (2014). N.Murak. & K.Iwats., Acta Phytotax. Geobot. 56(2): 127–128.
Monophyletic (Sundue et al., 2014). 12 species. 2005. Type: Claytosmunda claytoniana (L.) Metzgar & Rouhan
( Osmunda claytoniana L.).
Themelium (T.Moore) Parris, Kew Bull. 52(3): 737.
1997. Type: Themelium tenuisectum (Blume) Claytosmunda claytoniana (L.) Metzgar & Rouhan, comb. nov.
Parris ( Polypodium tenuisectum Blume). Cir- Osmunda claytoniana L., Sp. Pl. 2: 1066. 1753.
cumscription sensu Parris (1997, 2004, 2010b) and
Sundue et al. (2014). Themelium is monophyletic Dennstaedtiineae Schwartsb. & Hovenkamp, subord. nov.
in (Sundue et al., 2014), but nested within a clade Type: Dennstaedtia Bernh. (1801) [1800]. Type: Dennstaedtia-
containing polyphyletic Oreogrammitis and Radio- ceae—Polypodiales.
grammitis. The types of Oreogrammitis and Rhizome generally dorsiventral and long-creeping, covered
Themelium remain to be sampled, and thus we with hairs or proto-scales (true scales absent); sori marginal or
recommend maintaining these genera pending sub-marginal, generally protected by marginal pseudo-indusia
further research. 29 species. or by laminar true indusia (sometimes both are present),
rarely fully exindusiate.
Tomophyllum (E.Fourn.) Parris, Gard. Bull.
Singapore 58(2): 245. 2007. Lectotype (desig-
nated by Parris, Gard. Bull. Singapore 58(2): 245. Grammitidoideae Parris & Sundue, stat. nov. Grammitida-
2007): Tomophyllum subsecundodissectum (Zoll.) ceae Newman, Hist. Brit. Ferns: 7. 1840, as “Grammitideae”.
Parris ( Polypodium subsecundodissectum Zoll.). Type: Grammitis Sw. (1801) [1800]—Polypodiaceae.
Circumscription sensu Parris (2007) and Sundue
et al. (2014). Monophyletic (Sundue et al., 2014). Lindsaeineae Lehtonen & Tuomisto, subord. nov. Type:
42 species. Lindsaea Dryand. ex J.Sm. (1793), as Lindsaeaceae—
Polypodiales.
Xiphopterella Parris, Gard. Bull. Singapore 58(2): Rhizome short- to long-creeping (rarely ascending), cov-
249. 2007. Type: Xiphopterella hieronymusii (C. ered with non-clathrate scales or rarely hairs; petioles with
Chr.) Parris ( Polypodium hieronymusii C.Chr.). one or rarely two (or several fusing into two in the upper part
Circumscription sensu Parris (2007) and Sundue of stipe) vascular bundles; sori marginal to sub-marginal,
et al. (2014). Monophyletic (Sundue et al., 2014). generally protected by laminar true indusia or rarely by
11 species. marginal pseudo-indusia or both.
Zygophlebia L.E.Bishop, Amer. Fern J. 79(3): 107.

Mankyuoideae J.R.Grant & B.Dauphin, subfam. nov. Type:
1989. Type: Zygophlebia sectifrons (Kunze ex
Mankyua B.Y.Sun, M.H.Kim & C.H.Kim—Ophioglossaceae.
Mett.) L.E.Bishop ( Polypodium sectifrons Kunze
Based on the diagnosis associated with Mankyua B.Y.Sun,
ex Mett.). Circumscription sensu Bishop (1989)
M.H.Kim & C.H.Kim, Taxon 50(4): 1020, f. 1. 2002.
and Sundue et al. (2014). Paraphyletic relative to
Enterosora (Sundue et al., 2014). This could be
resolved by placing Zygophlebia in synonymy. Phegopteridoideae Salino, A.R.Sm. & T.E.Almeida, subfam.
However, this would be premature since the type e (1852)—Thelypteridaceae.
nov. Type: Phegopteris (C.Presl) Fe
of Enterosora has not yet been sampled and its Laminae bipinnate-pinnatifid or more divided, or laminae
placement is not certain. 20 species. bipinnatifid or tripinnatifid; costae adaxially lacking grooves;
veins free, not reaching the margins; chromosome base
Subfamily placement uncertain. numbers x ¼ 30 or 31.
Synammia C.Presl, Tent. Pterid.: 212, t.9, f.11. Pteridineae J.Prado & Schuettp., stat. nov. Pteridaceae E.D.
1836. Lectotype (designated by Copel., Ann. M.Kirchn., Schul-Bot. 109. 1831. Type: Pteris L. (1753)—
Cryptog. Phytopathol. 5 [Gen. Fil.]: 212. 1947): Polypodiales.
Synammia triloba (Cav.) C.Presl. Circumscription
sensu Schneider et al. (2006). Monophyletic Saccolomatineae Hovenkamp, subord. nov. Type: Saccoloma
(Schneider et al., 2006). Three species. Kaulf. (1820)—Saccolomataceae.
Rhizome erect, covered with multistratose, non-clathrate
scales, in cross-section with two concentric rings of mer-
Nomenclatural novelties isteles, sori cup-shaped, spores trilete, tetrahedral.
Aspleniineae H.Schneid. & C.J.Rothf., stat. nov. Asplenia-

ceae Newman, Hist. Brit. Ferns 6. 1840. Type: Asplenium L. Acknowledgements
(1753)—Polypodiales. We thank all pteridologists, past and present, who have
provided the taxonomic and phylogenetic foundation for this
Claytosmunda (Y.Yatabe, N.Murak. & K.Iwats.) Metzgar & effort. We are also grateful to Kanchi Gandhi for helping with
Rouhan, stat. nov. Osmunda subg. Claytosmunda Y.Yatabe, some nomenclatural aspects.

PPG I 595
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Journal of Systematics

JSE and Evolution doi: 10.1111/jse.12229

A community-derived classiﬁcation for extant lycophytes

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Austrolycopodium Holub, Folia Geobot. Phytotax.

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Dicranopteris Bernh., Neues J. Bot. 1(2): 38–39. Monophyletic (Wikstr€

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19. Family Loxsomataceae C.Presl, Gef€assb€

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sensu Tryon et al. (1990). Includes: Trismeria F

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v., Fl. Kouy-Tche

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S.K.Wu & K.L.Phan. Monophyletic (Kreier et al., Le

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Zygophlebia L.E.Bishop, Amer. Fern J. 79(3): 107.

Aspleniineae H.Schneid. & C.J.Rothf., stat. nov. Asplenia-

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J. Syst. Evol. 54 (6): 563–603, 2016 www.jse.ac.cn

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J. Syst. Evol. 54 (6): 563–603, 2016 www.jse.ac.cn

www.jse.ac.cn J. Syst. Evol. 54 (6): 563–603, 2016

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