JSE Journal of Systematics 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, Tha
ıs Elias Almeida9, Barbara Parris10, Emily B. Sessa11, Ashley R. Field12,
Lu
ıs de Gasper13, Carl J. Rothfels14, Michael D. Windham15, Marcus Lehnert16, Benjamin Dauphin17, Andre 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, Vin
ıcius Antonio de Oliveira Dittrich33, Yi-Fan Duan34, Jean-Yves Dubuisson35, Donald R. Farrar36, Susan Fawcett8,
es-Neto7, Jason R. Grant17, Amanda L. Grusz38, Jose Mar
ıa Gabriel y Gal
an37, Luiz Armando de Ara
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 46 47 48 49
rez , Hong-Mei Liu , Ngan Thi Lu , Esteban I. Meza-Torres , Xin-Yuan Miao45, Robbin Moran50, Melanie Link-Pe Claudine Massi Mynssen51, Nathalie Nagalingum52, Benjamin Øllgaard53, Alison M. Paul54, Jovani B. de S. Pereira55,
nica Ponce56, Tom A. Ranker57, Christian Schulz55, Wataru Shinohara58, Alexander Shmakov59, Leon R. Perrie24, Mo 1 Erin M. Sigel , Filipe Soares de Souza7, Lana da Silva Sylvestre60, Weston Testo8, Luz Amparo Triana-Moreno61, Chie Tsutsumi18, Hanna Tuomisto19, Iv
an 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 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 Herb
ario SP, Instituto de Bot^ anica, 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 Bot^anica, Universidade Federal de Minas Gerais 8 Pringle Herbarium, Department of Plant Biology, University of Vermont 9 Herb
ario 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 Neuch^atel 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 R
ıo Cuarto 2 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 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 Biolog
ıa 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 Bot
anica del Nordeste, Universidad Nacional del Nordeste, Consejo Nacional de Investigaciones Cient
ıficas 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 Bot
anica Darwinion, Consejo Nacional de Investigaciones Cient
ıficas 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 Bot^anica, Universidade Federal do Rio de Janeiro 61 Departamento de Ciencias Biol
ogicas, Universidad de Caldas 62 Departamento de Bot
anica, Universidad de Panam
a 63 Instituto de Biolog
ıa, 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 27 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 Free-sporing vascular plants comprise two distinct evolutionary lineages—lycophytes and ferns (e.g., Kenrick & Crane, 1997; Pryer et al., 2001)—that share a unique life cycle with independent gametophyte and sporophyte phases (e.g., Haufler et al., 2016). Classifications of these plants—often referred to collectively as pteridophytes despite their phylogenetic separation—have long been grounded in evolutionary principles and have reflected perceived evolutionary trends (e.g., Copeland, 1947; Pichi Sermolli, 1973, 1977b; Kramer & Green, 1990; Smith, 1995; Smith et al., 2006b; Christenhusz & Chase, 2014). However, these classifications were often conflicting, in large part due to a paucity of information concerning pteridophyte relationships and a lack of consensus regarding patterns of morphological evolution. The increased availability of molecular data and advances in phylogenetic methods have revolutionized our understanding of lycophyte and fern relationships. Today, as a consensus emerges concerning the evolutionary history of these lineages, classifications aimed at recognizing their natural groups are becoming increasingly “predictive” (Stuessy, 2009) and, we hope, stable. The ordinal and familial scheme by Smith et al. (2006b) was the first higher-level pteridophyte classification published in the molecular era. Although this work focused exclusively on ferns, these plants—also referred to as “monilophytes” in some publications (e.g., Pryer et al., 2004; Smith et al., 2006b; Schneider et al., 2009; here we simply use “ferns”)—account for nearly 90% of extant pteridophyte diversity. Founded on the principle of monophyly, while also recognizing the importance of maintaining well-established names and circumscriptions, the Smith et al. (2006b) classification established a new standard in fern taxonomy. Over the past decade, many important advances have been made in our understanding of relationships, some of which are reflected in subsequent higher-level schemes (e.g., Christenhusz et al., 2011; Rothfels et al., 2012b). A recent scheme by Christenhusz & Chase (2014) was also consistent with an understanding of fern phylogeny but represented a considerable departure in terms of stability and has not been widely adopted. Here, we provide a modern, comprehensive classification for lycophytes and ferns, following the main tenets of the Smith et al. (2006b) work but utilizing a democratic community-based approach. To accomplish this goal, we have established the Pteridophyte Phylogeny Group (PPG), based loosely on the model employed for flowering plant classification (APG, 1998; APG II, 2003; APG III, 2009; APG IV, 2016). Below, we outline our general philosophy and approach, summarize our outcomes, and present a consensus classification for all pteridophytes, from the rank of class to that of genus. Philosophy Biological classifications are fundamentally tools for communication 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 recognition and the ranks at which to treat them. A focus on natural groups is similarly important, as it results in classifications that reflect evolutionary history (Schmidt-Lebuhn, www.jse.ac.cn 565 2011) and ultimately lead to greater stability. Although we acknowledge the validity of some arguments presented in favor of maintaining paraphyletic taxa (see, e.g., H€ orandl & Stuessy, 2010), we ultimately reject this practice and aim to recognize only monophyletic lineages at the genus level and above. There are many reasons why monophyly might not be expected at the species level. This is especially true when polyploid speciation is rampant and, among pteridophytes, it has been estimated that nearly a third of all speciation events are correlated with an increase in ploidy (Wagner & Wagner, 1980; Wood et al., 2009). We acknowledge the difficulties inherent in translating evolutionary tree structures into hierarchical rank-based classifications. However, because such classifications have been implemented for centuries and continue to be used extensively, we do not subscribe to the alternative approach (disruptive, from the standpoint of stability) recommended in the PhyloCode (Cantino & de Queiroz, 2000). Due to the many challenges of incorporating fossil taxa, this first installment of the PPG classification focuses exclusively on extant lycophytes and ferns. Although some fossils could easily be accommodated, the phylogenetic affinities of most extinct plants are rather unclear. Many fossil taxa represent distinct evolutionary lineages and their inclusion in the PPG I classification would not only require revised circumscriptions, but almost certainly the recognition of new families, orders, and even classes. In general, the PPG I classification seeks to preserve existing taxa and circumscriptions that are widely accepted and consistent with our understanding of lycophyte and fern phylogeny. Although monophyly is our primary criterion for the recognition of taxa (Backlund & Bremer, 1998), we adopt a conservative approach that minimizes name changes in cases where data are insufficient to circumscribe what we judge to be stable monophyletic groups; this is with the understanding that some currently accepted taxa may ultimately be revealed to be non-monophyletic. In making decisions, we secondarily weigh morphological diagnosability and homogeneity, and hierarchical equivalency in terms of both age and diversity. This classification is not meant to be the final word on pteridophyte 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 recent literature on lycophyte and fern phylogeny, a framework for guiding future investigations, and a stimulus to further discourse. Although we welcome its use by systematists, other scientists, herbaria, governmental agencies, and others with a professional or non-professional interest in plants, we recognize that a single system may not 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, as noted below. Approach Establishment of the Pteridophyte Phylogeny Group (PPG) began through promotion of the concept at international conferences (including the 2015 Next Generation Pteridology J. Syst. Evol. 54 (6): 563–603, 2016 566 The Pteridophyte Phylogeny Group conference) and posts to society websites and e-mail lists, including those of the American Fern Society (AFS) and the International Association of Pteridologists (IAP). After a series of informal discussions among the early respondents, an electronic mailing list was established to enhance community engagement and facilitate discussion. As the project gained momentum, the community of collaborators expanded; currently, the PPG is a global community of 94 pteridologists. The classification presented herein is community-derived, and the taxa recognized have the support of a majority (or a plurality, in select instances) of the contributors. Decisions were made using an inclusive, transparent, and straightforward approach that typically involved: (1) an initial conversation among members of a focused subcommittee consisting of experts willing to contribute to the discussion; (2) formulation of a draft proposal by the subcommittee; (3) discussion of the draft proposal among all members of the 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 the whole of the PPG. After discussion of the advantages and disadvantages of various options, one system (recognizing classes and subclasses) emerged as the clear preference. A vote was taken to ensure there was majority support. A similar approach was used later to reach decisions concerning the informal names to be applied to major clades. To determine the orders, suborders, and families to be recognized, an organizing subcommittee developed the initial proposal. This was then shared electronically with all contributors, along with a survey developed to gauge acceptability and highlight areas needing further discussion. With the results of this survey in hand, discussion proceeded among all contributors. Following this dialogue, a revised poll was constructed that presented alternatives in areas of disagreement. All orders, suborders, and families recognized herein ultimately received majority support, ranging from just over 50% to unanimous approval. After settling on a list of higher-level taxa to be recognized, members of the PPG were asked (via an online survey) whether they felt that the generic classification within particular clades was in need of attention. In cases where a unanimous consensus existed, the organizers implemented the prevailing generic classification. If a clade were identified as needing attention, a subcommittee was established, consisting of contributors with experience in the particular lineage. Clade-focused subcommittees, under the leadership of subcommittee heads (selected by subcommittee members), discussed alternative generic classifications, reached a consensus, and presented a proposal for approval by the whole of the PPG. Any concerns were mediated via open dialogue through the PPG e-mail list. After arriving at a system of classes, subclasses, orders, suborders, families, and genera to be recognized, subcommittee heads (with the assistance of other contributors) were then tasked with compiling the following pertinent information for each included taxon: (1) the author of the name and the protologue reference; (2) for genera, the type species and basionym, along with the lectotypification reference (where applicable); (3) a brief statement outlining the circumscription J. Syst. Evol. 54 (6): 563–603, 2016 of the taxon; (4) a brief statement concerning its monophyly; (5) a list of any included taxa (i.e., synonyms); (6) the number of species attributed to the taxon; and (7) any additional comments. The classification presented herein reflects our current understanding of lycophyte and fern phylogeny (see references below). But, ultimately, circumscriptions adopted herein are all subject to reconsideration, given new evidence. Future research will enable greater insight into pteridophyte evolution and our classification will need to accommodate more robust hypotheses. To this end, we envision formal updates to the PPG I classification, with interim improvements tracked on a PPG website. Such adjustments (most likely a series of small incremental changes) will be carried out via the process outlined above, with initial proposals generated by smaller groups that are subsequently discussed and approved or rejected by the broader PPG. Outcomes We recognize two pteridophyte classes: Lycopodiopsida (lycophytes) and Polypodiopsida (ferns). These are distinct lineages within the tracheophyte tree of life, with ferns resolved as more closely related to seed plants than to lycophytes (Fig. 1; Kenrick & Crane, 1997; Pryer et al., 2001). Within Lycopodiopsida, we further recognize three orders (Lycopodiales, Isoëtales, and Selaginellales). Order Lycopodiales includes one family and 16 genera, whereas orders Isoëtales and Selaginellales each contain a single monogeneric family. Within Polypodiopsida, we recognize four subclasses: Equisetidae (horsetails); Ophioglossidae; Marattiidae; and Polypodiidae (leptosporangiates). Extant Equisetidae includes a single order, a single small family, and a single genus (Equisetum L.). Subclass Ophioglossidae encompasses two orders, each with a single family, and a total of 12 genera. Marattiidae includes just one order, one family, and six genera. Subclass Polypodiidae comprises the vast majority of extant fern diversity (Fig. 1). Here, we recognize seven orders (Osmundales, Hymenophyllales, Gleicheniales, Schizaeales, Salviniales, Cyatheales, and Polypodiales), with the Polypodiales subsequently divided into six suborders (Saccolomatineae, Lindsaeineae, Pteridineae, Dennstaedtiineae, Aspleniineae, and Polypodiineae). Within Osmundales, we recognize a single small family with six genera. Order Hymenophyllales also includes just one family, but it is considerably larger and encompasses nine genera. Gleicheniales and Schizaeales each constitute three relatively small families, with ten and four total genera, respectively. The order Salviniales comprises two families and five genera, and the order Cyatheales encompasses seven small families, with a total of ten genera, plus the larger Cyatheaceae with three genera. Within the Polypodiales, suborder Saccolomatineae includes a single small monogeneric family (Saccolomataceae). Suborder Lindsaeineae comprises two monogeneric families plus the larger Lindsaeaceae with seven genera. We recognize a single large family within Pteridineae, which includes 52 genera. Suborder Dennstaedtiineae also includes a single family, with ten genera. The remaining two suborders, www.jse.ac.cn 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. www.jse.ac.cn J. Syst. Evol. 54 (6): 563–603, 2016 568 The Pteridophyte Phylogeny Group Aspleniineae (eupolypods II) and Polypodiineae (eupolypods I), are exceedingly diverse, together accounting for well over half of extant pteridophyte diversity. Suborder Aspleniineae encompasses four large families, with a total of 59 genera, and seven smaller families, with 14 genera and fewer than 100 species. Finally, suborder Polypodiineae comprises three large families, with a total of 98 genera, and six small families, with ten genera. In all, this PPG I classification treats an estimated 11 916 species in 337 genera, 51 families, 14 orders, and 2 classes. In some cases, additional divisions are made to recognize subclasses and suborders (as noted above), as well as subfamilies (see classification below). Classes, subclasses, orders, suborders, and families, are organized phylogenetically, generally with increasing species diversity (Fig. 1). Within each family, subfamilies, if recognized, are organized by increasing species diversity; genera are then organized alphabetically within each family or subfamily, as applicable. Discussion As noted above, the PPG I classification presented here reflects our current, collective understanding of lycophyte and fern phylogeny. It takes into consideration all studies examining the relationships of these plants published through mid-2016. The PPG I classification also aims to maintain, where possible, continuity in the naming and circumscription of families and genera as recognized by most pteridologists prior to the molecular revolution (e.g., Kramer & Green, 1990). In this respect, this study continues the approach employed in most recent classifications (e.g., Smith et al., 2006b; Christenhusz et al., 2011; Rothfels et al., 2012b) but differs from classifications that favored a reduction in the number of taxa recognized (e.g., Christenhusz & Chase, 2014). At the highest taxonomic ranks, the PPG I classification is wholly compatible with a scheme recently proposed for all of life. Ruggiero et al. (2015) treated lycophytes and ferns as distinct subphyla (Lycopodiophytina and Polypodiophytina), each with a single class (Lycopodiopsida and Polypodiopsida). The present classification recognizes the two classes but does not make recommendations above this rank. Further, the PPG I classification agrees with Ruggiero et al. (2015) in recognizing four subclasses within ferns (Equisetidae, Ophioglossidae, Marattiidae, and Polypodiidae). This approach differs from that of Smith et al. (2006b), in which the four major lineages of ferns were recognized as classes, as well as from that of Chase & Reveal (2009), where embryophytes were treated collectively as a single class, but is consistent with the scheme used in the Tree of Life Web Project (Pryer et al., 2009). Although two of these higher-level classifications (Smith et al., 2006b; Chase & Reveal, 2009) do not provide a formal scientific name for ferns, they do both clearly unite the four fern lineages that we recognize as subclasses (Equisetidae, Ophioglossidae, Marattiidae, and Polypodiidae). We do not consider our decision to include them together in a single class to be controversial, as the corresponding clade (i.e., ferns) has been resolved consistently in analyses of molecular data (e.g., Pryer et al., 2001, 2004; Qiu et al., 2007; Schneider et al., 2009; Ruhfel et al., 2014; Wickett et al., 2014; Rothfels et al., 2015) and even J. Syst. Evol. 54 (6): 563–603, 2016 most analyses of morphological data (Kenrick & Crane, 1997; Schneider et al., 2009). Pteridophyte classification has been exceedingly stable at the ordinal level over the past decade. Smith et al. (2006b) recognized 11 orders within ferns, one for each of four major eusporangiate lineages and seven within leptosporangiates. Christenhusz et al. (2011) and Christenhusz & Chase (2014) adopted these same 11 fern orders plus three orders for lycophytes (not treated by Smith et al., 2006b). At the ordinal level, the PPG I classification does not differ at all from the earlier classifications, but we do introduce a new informal name for one large order. Polypodiales, which accounts for more than 80% of fern diversity, has previously been referred to as “polypods” (Schneider et al., 2004b; Smith et al., 2006b). However, “polypods” has also been used to refer to the members of the family Polypodiaceae that do not belong to the distinct (and diverse) “grammitid” clade (Smith, 1995; Sundue et al., 2014). Here, we aim to avoid confusion by using 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 annular cells—that characterizes this lineage. Within the Polypodiales, we recognize six suborders, providing formal names for “eupolypods I”, “eupolypods II”, and other clades. As this rank has not typically been utilized within ferns, five of these suborders are new. Outside of the Polypodiales, the 25 families recognized in this PPG I classification are identical to those in Smith et al. (2006b, for ferns) and Christenhusz et al. (2011). Within the Polypodiales, we recognize 26 families, and there are some stark differences relative to Smith et al. (2006b) and Christenhusz et al. (2011). The Lindsaeaceae, Woodsiaceae, Dryopteridaceae, and Lomariopsidaceae sensu Smith et al. (2006b) have all been found to be paraphyletic in more recent phylogenetic analyses and are here divided into three, seven, three, and two families, respectively. The present PPG I classification additionally recognizes subfamilies within some of the larger families of lycophytes and ferns, especially those with a history of subdivision. This has the advantage of providing formal names for some clades (e.g., vittarioids and grammitids) that were long recognized as distinct families but were later subsumed into other families (within which they appear to be nested). Unlike recent comprehensive treatments of lycophytes and ferns (e.g., Smith et al., 2006b; Christenhusz et al., 2011), the PPG I classification extends to the genus level. We formally recognize 337 pteridophyte genera, representing a 50% increase over the 223 genera included in the pre-molecular system of Kramer & Green (1990). Although this difference is dramatic, the constituent changes have, in fact, accumulated incrementally over the past quarter-century. When existing generic concepts are found to be in conflict with the results of molecular phylogenetic analyses, there are, using monophyly as a criterion, two paths to resolution: disintegration (i.e., the “splitting” of a non-monophyletic genus into multiple monophyletic genera to maintain a nested genus); or integration (i.e., the “lumping” of a nested genus with the genus it renders non-monophyletic). Both approaches have an impact on stability, as the former results in new descriptions (or reinstatements) and the latter in synonymizations. www.jse.ac.cn PPG I Looking again at the marked increase in the number of pteridophyte genera, it is clear that disintegration has typically been favored and some lineages have experienced greater change than others. Instead of recognizing just four genera within the Lycopodiaceae (sensu Øllgaard, 1990), the PPG I classification takes the now-common approach of subdividing the family into 16 genera (Holub, 1964, 1975, 1983, 1985, 1991; Wagner & Beitel, 1992; Haines, 2003; Øllgaard, 2012a, 2012b, 2015; Field & Bostock, 2013; Øllgaard & Windisch, 2014; Field et al., 2016). Likewise, we mostly follow Almeida et al. (2016) in our treatment of 30 genera in Thelypteridaceae (versus five in Smith, 1990) and we also incorporate considerable subdivision recently proposed for the Hymenophyllaceae (sensu Ebihara et al., 2006), grammitids (sensu Sundue et al., 2014), and Blechnaceae (sensu Gasper et al., 2016b). In the majority of these studies, the authors were largely able to re-establish preexisting (narrower) concepts that were both introduced and discarded prior to the molecular revolution. Of course, more inclusive generic concepts have been favored in some cases. We continue to recognize a broad Asplenium L. (segregating only Hymenasplenium Hayata, sensu Schneider et al., 2004a). And, despite recent subdivisions, the present PPG I classification ultimately recognizes an inclusive Davallia Sm. (sensu Tsutsumi et al., 2016) and Woodsia R.Br. (sensu Shao et al., 2015). Furthermore, the genus Dryopteris Adans. is greatly expanded here, relative to its definition at the end of the 20th century (Kramer et al., 1990), to include segregates such as Acrophorus C.Presl, Dryopsis Holttum & P.J.Edwards, Nothoperanema (Tagawa) Ching, and Peranema D.Don (Zhang & Zhang, 2012; Zhang et al., 2012). Notably, resolution via integration, and the resulting recognition of larger genera, appears to be much more common in flowering plants (Humphreys & Linder, 2009). The PPG I classification presented below does not provide species lists; however, we do present estimates of species number for each genus. Interestingly, our genus-based totals (1338 lycophyte species and 10 578 fern species) are remarkably similar to the family-based sums provided by Christenhusz & Chase (2014; 1300 lycophytes and 10 535 ferns). Classification Class Lycopodiopsida Bartl., Ord. Nat. Pl.: 14, 19. 1830. Circumscription sensu Ruggiero et al. (2015). Monophyletic (Pryer et al., 2001; Kenrick & Crane, 1997; Rai & Graham, 2010; Wickett et al., 2014). Three orders, three families, 18 genera, and an estimated 1338 species. A. Order Lycopodiales DC. ex Bercht. & J.Presl, Prir. Rostlin: 272. 1820. Circumscription equivalent to family Lycopo€m & diaceae in this classification. Monophyletic (Wikstro Kenrick, 1997, 2000, 2001; Ji et al., 2008; Field et al., 2016). One family, 16 genera, and an estimated 388 species. 1. Family Lycopodiaceae P.Beauv. in Mirb., Hist. Nat. Veg. 4: 293. 1802. Circumscription sensu Øllgaard (2015). €m & Kenrick, 1997, 2000, 2001; Monophyletic (Wikstro www.jse.ac.cn 569 Ji et al., 2008; Field et al., 2016). Three subfamilies, 16 genera, and an estimated 388 species. Subfamily Lycopodielloideae W.H.Wagner & Beitel ex B.Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription sensu Øllgaard (2015). Monophyletic (Field et al., 2016). Included in Lycopodioideae in Field et al. (2016). Four genera and an estimated 54 species. Lateristachys Holub, Folia Geobot. Phytotax. 18: 440. 1983. Type: Lateristachys lateralis (R.Br.) Holub ( Lycopodium laterale R.Br.). Circumscription sensu Holub (1983). Monophyletic (Field et al., 2016). Four species. Lycopodiella Holub, Preslia 36: 20, 22. 1964. Type: Lycopodiella inundata (L.) Holub ( Lycopodium inundatum L.). Circumscription sensu Øllgaard & Windisch (2014). Monophyletic (Field et al., 2016). 15 species. Palhinhaea Franco & Vasc., Bol. Soc. Brot., ser. 2, 41: 24. 1967. Type: Palhinhaea cernua (L.) Vasc. & Franco ( Lycopodium cernuum L.). Circumscription sensu Holub (1985, 1991), Øllgaard (2012b), and Øllgaard & Windisch (2014). Monophyletic (Field et al., 2016). Palhinhaea Franco & Vasc. is proposed to be conserved against Lepidotis P. Beauv. (Greuter & Troia, 2014). 25 species. Pseudolycopodiella Holub, Folia Geobot. Phytotax. 18: 441. 1983. Type: Pseudolycopodiella caroliniana (L.) Holub ( Lycopodium carolinianum L.). Circumscription sensu Holub (1983). Monophyletic (Field et al., 2016). 10 species. Subfamily Lycopodioideae W.H.Wagner & Beitel ex B. Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription sensu Øllgaard (2015). Monophyletic (Field et al., 2016). Including Lycopodielloideae in Field et al. (2016). Nine genera and an estimated 58 species. Austrolycopodium Holub, Folia Geobot. Phytotax. 26(1): 90. 1991. Type: Austrolycopodium magellanicum (P.Beauv.) Holub ( Lepidotis magellanica P. Beauv.). Circumscription sensu Holub (1991). Monophyletic (Field et al., 2016). Eight species. Dendrolycopodium A.Haines, Fam. Huperziac. Lycopodiac. New England: 84. 2003. Type: Dendrolycopodium obscurum (L.) A.Haines ( Lycopodium obscurum L.). Circumscription sensu Haines (2003). Monophyletic (Field et al., 2016). Four species. Diphasiastrum Holub, Preslia 47(2): 104. 1975. Type: Diphasiastrum complanatum (L.) Holub ( Lycopodium complanatum L.). Circumscription sensu Holub (1975, 1985). Monophyletic (Field et al., 2016). 20 species. J. Syst. Evol. 54 (6): 563–603, 2016 570 The Pteridophyte Phylogeny Group Diphasium C.Presl ex Rothm., Repert. Spec. Nov. Regni Veg. 54: 64. 1944. Type: Diphasium jussiaei (Desv.) Rothm. ( Lycopodium jussiaei Desv.). Circumscription sensu Holub (1985). Monophyletic (Field et al., 2016). Five species. Lycopodiastrum Holub ex R.D.Dixit, J. Bombay Nat. Hist. Soc. 77(3): 540. 1981. Type: Lycopodiastrum casuarinoides (Spring) Holub ex R.D.Dixit ( Lycopodium casuarinoides Spring). Circumscription sensu Holub (1983) and Zhang & Iwatsuki (2013). Monotypic (Field et al., 2016). Lycopodium L., Sp. Pl. 2: 1100. 1753. Lectotype (designated by W.J.Rob., Bull. Torrey Bot. Club 41: 51. 1914): Lycopodium clavatum L. Circumscription sensu Haines (2003). Monophyletic (Field et al., 2016). 15 species. Pseudodiphasium Holub, Folia Geobot. Phytotax. 18: 440. 1983. Type: Pseudodiphasium volubile (G. Forst.) Holub ( Lycopodium volubile G.Forst.). Circumscription sensu Holub (1983). Monotypic (Field et al., 2016). Pseudolycopodium Holub, Folia Geobot. Phytotax. 18: 441. 1983. Type: Pseudolycopodium densum (Rothm.) Holub ( Lepidotis densa Rothm.). Circumscription sensu Holub (1983). Monotypic (Field et al., 2016). Spinulum A.Haines, Fam. Huperziac. Lycopodiac. New England: 85. 2003. Type: Spinulum annotinum (L.) A.Haines ( Lycopodium annotinum L.). Circumscription sensu Haines (2003). Presumably monophyletic; only one species sampled to date (Field et al., 2016). Three species. Subfamily Huperzioideae W.H.Wagner & Beitel ex B. Øllg., Nordic J. Bot. 33(2): 195. 2015. Circumscription sensu Øllgaard (2015) and Field et al. (2016). Monophyletic (Field et al., 2016). Three genera and an estimated 276 species. Huperzia Bernh., J. Bot. (Schrader) 1800(2): 126. 1801. Lectotype (designated by Rothm., Repert. Spec. Nov. Regni Veg. 54: 59. 1944): Huperzia selago (L.) Bernh. ex Schrank & Mart. ( Lycopodium selago L.). Circumscription sensu Wagner & Beitel (1992), Zhang & Kung (1998, 2000a), Zhang & Iwatsuki (2013), Øllgaard (2015), and Field et al. (2016). Monophyletic (Field et al., 2016). 25 species. Phlegmariurus Holub, Preslia 36(1): 17, 21. 1964. Type: Phlegmariurus phlegmaria (L.) T.Sen & U.Sen ( Lycopodium phlegmaria L.). Circumscription sensu Wagner & Beitel (1992), Zhang & Kung (1999, 2000b), Øllgaard (2012a, 2012b, 2015), Field & Bostock (2013), Zhang & Iwatsuki (2013), Øllgaard & Windisch (2014), and Field et al. (2016). Monophyletic (Field et al., 2016). About 250 species. J. Syst. Evol. 54 (6): 563–603, 2016 Phylloglossum Kunze, Bot. Zeitung (Berlin) 1: 721. 1843. Type: Phylloglossum drummondii Kunze. Circumscription sensu Field et al. (2016). Monotypic (Field et al., 2016). B. Order Isoëtales Prantl, Lehrb. Bot.: 116, 125. 1874. Circumscription equivalent to family Isoëtaceae in this classification. Monophyletic (Rydin & Wikstr€ om, 2002;
n & Rydin, 2016). One family, one Hoot et al., 2006; Larse genus, and about 250 species. 2. Family Isoëtaceae Dumort., Anal. Fam. Pl.: 67. 1829. Circumscription sensu Jermy (1990a). Monophyletic €m, 2002; Hoot et al., 2006; Larse
n & (Rydin & Wikstro Rydin, 2016). One genus and about 250 species. Isoëtes L., Sp. Pl. 2: 1100. 1753. Type: Isoëtes lacustris L. Circumscription sensu Jermy (1990a). Monophy
n & Rydin, 2016). About 250 species. letic (Larse C. Order Selaginellales Prantl, Lehrb. Bot.: 116, 124. 1874. Circumscription equivalent to family Selaginellaceae in this classification. Monophyletic (Korall et al., 1999; Korall & Kenrick, 2004; Zhou et al., 2015; Weststrand & Korall, 2016a). One family, one genus, and perhaps 700 species. 3. Family Selaginellaceae Willk., Anleit. Stud. Bot. 2: 163. 1854. Circumscription sensu Jermy (1990b). Monophyletic (Korall et al., 1999; Korall & Kenrick, 2004; Zhou et al., 2015; Weststrand & Korall, 2016a). One genus and perhaps 700 species. Selaginella P.Beauv., Mag. Encycl. 9(5): 478. 1804. Type: Selaginella selaginoides (L.) P.Beauv. ex Schrank & Mart. ( Lycopodium selaginoides L.). Circumscription sensu Jermy (1990b). Monophyletic (Korall et al., 1999; Korall & Kenrick, 2004; Zhou et al., 2015; Weststrand & Korall, 2016a). Six (Zhou & Zhang, 2015) or seven (Weststrand & Korall, 2016b) subgenera can be recognized. Perhaps 700 species. Class Polypodiopsida Cronquist, Takht. & W.Zimm., Taxon 15: 133. 1966. Circumscription sensu Pryer et al. (2009) and Ruggiero et al. (2015). Monophyletic (Pryer et al., 2001, 2004; Qiu et al., 2007; Schneider et al., 2009; Ruhfel et al., 2014; Wickett et al., 2014; Rothfels et al., 2015). Four subclasses, 11 orders, 48 families, 319 genera, and an estimated 10 578 species. Subclass Equisetidae Warm., Osnov. Bot.: 221. 1883. Circumscription sensu Pryer et al. (2009) and Ruggiero et al. (2015). Subclass consists of one extant order, one family, one genus, and 15 species. D. Order Equisetales DC. ex Bercht. & J.Presl, Prir. Rostlin: 271. 1820. Circumscription sensu Smith et al. (2006b). Order consists of one extant family, one genus, and 15 species.
d. 4. Family Equisetaceae Michx. ex DC., Essai Propr. Me Pl.: 49. 1804. Circumscription sensu Smith et al. (2006b). Family consists of one extant genus and 15 species. www.jse.ac.cn PPG I Equisetum L., Sp. Pl. 2: 1061. 1753. Lectotype (designated by Hauke, Nova Hedwigia 30: 414. 1978): Equisetum fluviatile L. Circumscription sensu Hauke (1990). Monophyletic (Guillon, 2007). An isolated genus of closely related and recently diverged species (Des Marais et al., 2003). 15 species. Subclass Ophioglossidae Klinge, Fl. Est.-Liv-Churland 1: 94. 1882. Circumscription sensu Ruggiero et al. (2015) and equivalent to Psilotidae in Pryer et al. (2009). Monophyletic (Pryer et al. 2001, 2004; Wickett et al., 2014; Rothfels et al., 2015). Two orders, two families, 12 genera, and an estimated 129 species. E. Order Psilotales Prantl, Lehrb. Bot., ed. 5: 183. 1884. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004; Rothfels et al., 2015). One family, two genera, and an estimated 17 species. 5. Family Psilotaceae J.W.Griff. & Henfr., Microgr. Dict.: 540. 1855. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004). Two genera and an estimated 17 species. Psilotum Sw., J. Bot. (Schrader) 1800(2): 8, 109. 1801. Lectotype (designated by Brongn., Dict. Class. Hist. Nat. 9: 558. 1826): Psilotum triquetrum Sw. Circumscription sensu Kramer (1990a). Only one species sampled to date, but assumed to be monophyletic. Two species. Tmesipteris Bernh., J. Bot. (Schrader) 1800(2): 8, 131. 1801. Type: Tmesipteris tannensis (Spreng.) Bernh. ( Lycopodium tannense Spreng.). Circumscription sensu Kramer (1990a). Monophyletic (Perrie et al., 2010). About 15 species. F. Order Ophioglossales Link, Hort. Berol. 2: 151.1833. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004; Rothfels et al., 2015). One family, ten genera, and an estimated 112 species. 6. Family Ophioglossaceae Martinov, Tekhno-Bot. Slovar: 438. 1820. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004; Rothfels et al., 2015). Four subfamilies, ten genera, and an estimated 112 species. Subfamily Helminthostachyoideae C.Presl, Suppl. Tent. Pterid.: 58. 1845. Circumscription sensu Shinohara et al. (2013). Subfamily consists of a single monotypic genus. Helminthostachys Kaulf., Enum. Filic.: 28. 1824. Type: Helminthostachys zeylanica (L.) Hook. ( Osmunda zeylanica L.). Circumscription sensu Wagner (1990). Includes: Ophiala Desv. Monotypic (Hauk et al., 2003; Shinohara et al., 2013). Subfamily Mankyuoideae J.R.Grant & B.Dauphin, this classification (see below). Circumscription sensu Shinohara et al. (2013). Subfamily consists of a single monotypic genus. www.jse.ac.cn 571 Mankyua B.Y.Sun, M.H.Kim & C.H.Kim, Taxon 50 (4): 1020. 2002. Type: Mankyua chejuensis B.Y.Sun, M.H.Kim & C.H.Kim. Circumscription sensu Sun et al. (2009). Monotypic (Shinohara et al., 2013). Subfamily Ophioglossoideae C.Presl, Suppl. Tent. Pterid.: 47. 1845. Circumscription sensu Hauk et al. (2003). Monophyletic (Hauk et al., 2003; Shinohara et al., 2013). Four genera and an estimated 47 species. Cheiroglossa C.Presl, Suppl. Tent. Pterid.: 56. 1845. Type: Cheiroglossa palmata (L.) C.Presl ( Ophioglossum palmatum L.). Circumscription sensu Hauk et al. (2003). Monophyletic (Hauk et al., 2003; Shinohara et al., 2013). Two species. Ophioderma (Blume) Endl., Gen. Pl.: 66. 1836. Type: Ophioderma pendulum (L.) C.Presl ( Ophioglossum pendulum L.). Circumscription sensu Hauk et al. (2003). Monophyletic (Hauk et al., 2003; Shinohara et al., 2013). Three species. Ophioglossum L., Sp. Pl. 2: 1062. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 369. 1875): Ophioglossum vulgatum L. Circumscription sensu Pichi Sermolli (1977b). Monophyletic (Hauk et al., 2003; Shinohara et al., 2013). 41 species. Rhizoglossum C.Presl, Suppl. Tent. Pterid.: 47. 1845. Type: Rhizoglossum bergianum (Schltdl.) C.Presl ( Ophioglossum bergianum Schltdl.). Circumscription sensu Clausen (1938) and Pichi Sermolli (1977b). Monotypic. Subfamily Botrychioideae C.Presl, Suppl. Tent. Pterid.: 42. 1845. Circumscription sensu Hauk et al. (2003). Monophyletic (Hauk et al., 2003; Shinohara et al., 2013). Four genera and an estimated 63 species. Botrychium Sw., J. Bot. (Schrader) 1800(2): 110. 1801. Lectotype (designated by J.Sm., Hist. Fil.: 369. 1875): Botrychium lunaria (L.) Sw. ( Osmunda lunaria L.). Circumscription sensu Wagner (1990) and Dauphin et al. (2014). Monophyletic (Hauk et al., 2003). About 35 species. Botrypus Michx., Fl. Bor.-Amer. (Michaux) 2: 274. 1803. Lectotype (designated by Pic.Serm., Webbia 26: 497. 1972): Botrypus virginianus (L.) Michx. ( Osmunda virginiana L.). Circumscription sensu Hauk et al. (2003). Includes: Osmundopteris (Milde) Small. Not monophyletic (Hauk et al., 2003; Shinohara et al., 2013). About two species. Japanobotrychium Masam., Journ. Soc. Trop. Agric. Formosa 3: 246. 1931. Type: Japanobotrychium arisanense Masam. Monotypic (Shinohara et al., 2013). Sceptridium Lyon, Bot. Gaz. 40(6): 457. 1905. Lectotype (designated by R.T.Clausen, Mem. J. Syst. Evol. 54 (6): 563–603, 2016 572 The Pteridophyte Phylogeny Group Torrey Bot. Club 19(2): 24. 1938): Sceptridium obliquum (Muhl. ex Willd.) Lyon ( Botrychium obliquum Muhl. ex Willd.). Circumscription sensu Hauk et al. (2003). Monophyletic (Hauk et al., 2003). About 25 species. Subclass Polypodiidae Cronquist, Takht. & W.Zimm., Taxon 15: 133. 1966. Circumscription sensu Pryer et al. (2008) and Ruggiero et al. (2015). Monophyletic (Pryer et al., 2004; Rothfels et al., 2015). Seven orders, 44 families, 300 genera, and an estimated 10 323 species. Subclass Marattiidae Klinge, Fl. Est.-Liv-Churland 1: 93. 1882. Circumscription sensu Christenhusz (2009) and Ruggiero et al. (2015). Monophyletic (Pryer et al., 2004). One order, one family, six genera, and an estimated 111 species. H. Order Osmundales Link, Hort. Berol.: 445. 1833. Circumscription sensu Smith et al. (2006b). Order consists of one family. One family, six genera, and an estimated 18 species. 8. Family Osmundaceae Martinov, Tekhno-Bot. Slovar.: 445. 1820. Circumscription sensu Smith et al. (2006b). Monophyletic (Yatabe et al., 1999; Schuettpelz & Pryer, 2007; Metzgar et al., 2008). Six genera and an estimated 18 species. G. Order Marattiales Link, Hort. Berol. 2: 148. 1833. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004). One family, six genera, and an estimated 111 species. 7. Family Marattiaceae Kaulf., Enum. Filic.: 31. 1824. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004). Six genera and an estimated 111 species. Claytosmunda (Y.Yatabe, N.Murak. & K.Iwats.) Metzgar & Rouhan, this classification (see below). Type: Claytosmunda claytoniana (L.) Metzgar & Rouhan ( Osmunda claytoniana L.). Circumscription equivalent to Osmunda subgenus Claytosmunda in Yatabe et al. (2005). Monotypic. Angiopteris Hoffm., Commentat. Soc. Regiae Sci. Gott. 12: 29. 1796. Type: Angiopteris evecta (G.Forst.) Hoffm. ( Polypodium evectum G.Forst.). Circumscription sensu Murdock (2008). Includes: Archangiopteris Christ & Giesenh.; Clementea Cav.; Macroglossum Copel.; Protangiopteris Hayata; Protomarattia Hayata; Psilodochea C.Presl. Monophyletic (Murdock, 2008). Perhaps 30 species. Leptopteris C.Presl, Suppl. Tent. Pterid.: 70. 1845. Lectotype (designated by C.Chr., Index Filic. lvi.1906): Leptopteris hymenophylloides (A.Rich.) C.Presl ( Todea hymenophylloides A.Rich.). Circumscription sensu Kramer (1990o). Monophyletic (Yatabe et al., 1999; Metzgar et al., 2008; Bomfleur et al., 2015). Six species. Christensenia Maxon, Proc. Biol. Soc. Washington 18 (50): 239. 1905. Type: Christensenia aesculifolia (Blume) Maxon ( Aspidium aesculifolium Blume). Circumscription sensu Murdock (2008). Includes: Kaulfussia Blume.; Macrostoma Griff. Only one species sampled to date, but assumed to be monophyletic (Murdock, 2008). Two species. Osmunda L., Sp. Pl. 2: 1063. 1753. Lectotype (designated by L
eman, Dict. Sci. Nat. 37: 9. 1825): Osmunda regalis L. Circumscription equivalent to subgenus Osmunda in Kramer (1990o). Monophyletic (Yatabe et al., 1999; Metzgar et al., 2008; Bomfleur et al., 2015). About four species. Osmundastrum C.Presl, Gef€assb€ undel Farrn: 18. 1847. Type: Osmundastrum cinnamomeum (L.) C. Presl ( Osmunda cinnamomea L.). Circumscription sensu Yatabe et al. (2005). Monotypic (Miller, 1967; Yatabe et al., 1999; Metzgar et al., 2008). Danaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 420, pl.9, f.11. 1793. Type: Danaea nodosa (L.) Sm. ( Acrostichum nodosum L.). Circumscription sensu Murdock (2008). Includes: Heterodanaea C.Presl. 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. 1906): Plenasium banksiaefolium (C.Presl) C.Presl. ( Nephrodium banksiaefolium C.Presl). Circumscription equivalent to subgenus Plenasium in Kramer (1990o). Monophyletic (Yatabe et al., 1999; Metzgar et al., 2008; Bomfleur et al., 2015). About four species. Eupodium J.Sm. in Hooker, Gen. Fil. (Hooker): t.118. 1842. Type: Eupodium kaulfussii (J.Sm.) J.Sm. ( Marattia kaulfussii J.Sm.). Circumscription sensu Murdock (2008). Monophyletic (Murdock, 2008). Two species. Marattia Sw., Prodr. 8: 128. 1788. Type: Marattia alata Sw. Circumscription sensu Murdock (2008). Includes: Gymnotheca C.Presl; Stibasia C.Presl. Monophyletic (Murdock, 2008). Seven species. Ptisana Murdock, Taxon 57(3): 744. 2008. Type: Ptisana salicina (Sm.) Murdock ( Marattia salicina Sm.). Circumscription sensu Murdock (2008). Monophyletic (Murdock, 2008). 20 species. J. Syst. Evol. 54 (6): 563–603, 2016 Todea Willd. ex Bernh., J. Bot. (Schrader) 1800(2): 126. 1801. Type: Todea africana Willd. ex Bernh., nom illeg. superfl. ( Acrostichum barbarum L.). Circumscription sensu Kramer (1990o). Monophyletic (Metzgar et al., 2008). Two species. I. Order Hymenophyllales A.B.Frank, Syn. Pflanzenk., ed. 2, 3: 1452. 1877. Circumscription sensu Smith et al. (2006b). www.jse.ac.cn PPG I Order consists of one family, nine genera, and an estimated 434 species. 9. Family Hymenophyllaceae Mart., Consp. Regn. Veg.: 3. 1835. Circumscription sensu Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007). Two subfamilies, nine genera, and an estimated 434 species. Subfamily Trichomanoideae C.Presl, Hymenophyllaceae: 102. 1843. Circumscription equivalent to “trichomanoids” in Ebihara et al. (2006). Monophyletic (Pryer et al., 2001; Dubuisson et al., 2003; Ebihara et al., 2007). Relationships among the trichomanoid genera are still mostly unresolved. Eight genera and an estimated 184 species. Abrodictyum C.Presl, Hymenophyllaceae: 20. 1843. Type: Abrodictyum cumingii C.Presl. Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). About 25 species. 573 Trichomanes radicans Sw.). Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). About 15 species. Subfamily Hymenophylloideae Burnett, Outlines Bot.: 324. 1835. Circumscription equivalent to “hymenophylloids” in Ebihara et al. (2006). Subfamily consists of one genus and an estimated 250 species.
m. Acad. Roy. Sci. (Turin) Hymenophyllum Sm., Me 5: 418, pl.9, f.8. 1793. Lectotype (designated by C.Presl, Hymenophyllaceae: 1. 1844): Hymenophyllum tunbrigense (L.) Sm. ( Trichomanes tunbrigense L.). Circumscription sensu Ebihara et al. (2006). Monophyletic (Pryer et al., 2001; Hennequin et al., 2006; Ebihara et al., 2007). About 250 species.
Pale
ont. Ve
g. 1: 669. J. Order Gleicheniales Schimp., Traite 1869. Circumscription sensu Smith et al. (2006b). Monophyletic (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. ( Trichomanes apiifolium C.Presl). Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). Five species. 10. Family Matoniaceae C.Presl, Gef€assb€ undel Farrn: 32. 1847. Circumscription sensu Smith et al. (2006b). Monophyletic (Kato & Setoguchi, 1998). Two genera and an estimated four species. Cephalomanes C.Presl, Hymenophyllaceae: 17. 1843. Type: Cephalomanes atrovirens C.Presl. Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). Four species. Matonia R.Br. ex Wall., Pl. Asiat. Rar. 1(1): 16. 1829. Type: Matonia pectinata R.Br. Circumscription sensu Kramer (1990l). Monophyletic (Kato & Setoguchi, 1998). Two species. Crepidomanes C.Presl, Epimel. Bot.: 258. 1851. Type: Crepidomanes intramarginale (Hook. & Grev.) C.Presl ( Trichomanes intramarginale Hook. & Grev.). Circumscription sensu Ebihara et al. (2006). Includes: Nesopteris Copel. Monophyletic (Ebihara et al., 2007). About 30 species. Phanerosorus Copel., Philipp. J. Sci. C 3(6): 344. 1909. Type: Phanerosorus sarmentosus (Baker) Copel. ( Matonia sarmentosa Baker). Circumscription sensu Kramer (1990l). Monophyletic (Kato & Setoguchi, 1998). Two species. Didymoglossum Desv., M
em. Soc. Linn. Paris 6: 330. 1827. Lectotype (designated by C.Chr., Index Filic. xiv. 1906): Didymoglossum hymenoides (Hedw.) Copel. ( Trichomanes hymenoides Hedw.). Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). About 30 species. Polyphlebium Copel., Philipp. J. Sci. 67: 55. 1938. Type: Polyphlebium venosum (R.Br.) Copel. ( Trichomanes venosum R.Br.). Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). About 15 species. Trichomanes L., Sp. Pl. 2: 1097. 1753. Lectotype (designated by Underw., Mem. Torrey Bot. Club 6: 256, 283. 1899): Trichomanes crispum L. Circumscription sensu Ebihara et al. (2006). Monophyletic (Ebihara et al., 2007). About 60 species. Vandenboschia Copel., Philipp. J. Sci. 67: 51. 1938. Type: Vandenboschia radicans (Sw.) Copel. ( www.jse.ac.cn 11. Family Dipteridaceae Seward & E.Dale, Philos. Trans., Ser. B. 194: 487, 499, 502. 1901. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004). Two genera and an estimated 11 species. Cheiropleuria C.Presl, Epimel. Bot.: 189. 1851. Lectotype (designated by J.Sm., Hist. Fil.: 139. 1875): Cheiropleuria bicuspis (Blume) C.Presl ( Polypodium bicuspe Blume). Circumscription sensu Kramer (1990b). Monophyletic (Kato et al., 2001). Three species. Dipteris Reinw., Syll. Pl. Nov. 2: 3. 1825. Type: Dipteris conjugata Reinw. Circumscription sensu Kramer (1990f). Presumably monophyletic (sampling limited to date). About eight species. 12. Family Gleicheniaceae C.Presl, Reliq. Haenk. 1: 70. 1825. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 2004). Six genera and an estimated 157 species. J. Syst. Evol. 54 (6): 563–603, 2016 574 The Pteridophyte Phylogeny Group Dicranopteris Bernh., Neues J. Bot. 1(2): 38–39. 1805. Type: Dicranopteris dichotoma (Thunb.) Bernh. ( Polypodium dichotomum Thunb.). Circumscription sensu Kramer (1990g), but excluding Gleichenella. Sampling to date has been insufficient to fully assess the monophyly of the genus (Li et al., 2010). Perhaps 20 species. Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 47. 1950. Lectotype (designated by C. Chr., Index Filic. liv. 1906): Diplopterygium glaucum (Thunb. ex Houtt.) Nakai ( Polypodium glaucum Thunb. ex Houtt.). Circumscription sensu Kramer (1990g). Sampling to date has been insufficient to fully assess the monophyly of the genus (Li et al., 2010). About 25 species. Gleichenella Ching, Sunyatsenia 5: 276. 1940. Type: Gleichenella pectinata (Willd.) Ching ( Mertensia pectinata Willd.). Circumscription sensu Mickel & Smith (2004). Monotypic.
m. Acad. Roy. Sci. (Turin) 5: Gleichenia Sm., Me 419, pl.9, f.10. 1793. Type: Gleichenia polypodioides (L.) Sm. ( Onoclea polypodioides L.). Circumscription sensu Kramer (1990g). Apparently monophyletic (Perrie et al., 2007; but see Hennequin et al., 2014), although sampling to date has been limited. About 15 species. Sticherus C.Presl, Tent. Pterid.: 51. 1836. Lectotype (designated by C.Chr., Index Filic. liv. 1906): Sticherus laevigatus (Willd.) C.Presl ( Mertensia laevigata Willd.). Circumscription sensu Kramer (1990g). Apparently monophyletic (Perrie et al., 2007), although sampling to date has been limited. About 95 species. Stromatopteris Mett., Ann. Sci. Nat. Bot., ser. 4, 15: 84. 1861. Type: Stromatopteris moniliformis Mett. Circumscription sensu Kramer (1990g). Monotypic.
Pale
ont. Ve
g. 1: 674. K. Order Schizaeales Schimp., Traite 1869. Circumscription sensu Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007). Three families, four genera, and an estimated 190 species. 13. Family Lygodiaceae M.Roem., Handb. Allg. Bot. 3: 520. 1840. Circumscription sensu Smith et al. (2006b). € m et al., 2002). One genus and Monophyletic (Wikstro an estimated 40 species. Lygodium Sw., J. Bot. (Schrader) 1800(2): 7, 106. 1801. Type: Lygodium scandens (L.) Sw. ( Ophioglossum scandens L.). Circumscription sensu €m et al., Kramer (1990q). Monophyletic (Wikstro 2002). Perhaps 40 species. 14. Family Schizaeaceae Kaulf., Wesen Farrenkr.: 119. 1827. Circumscription sensu Smith et al. (2006b). J. Syst. Evol. 54 (6): 563–603, 2016 Monophyletic (Wikstr€ om et al., 2002). Two genera and an estimated 35 species. Actinostachys Wall., Numer. List 1. 1829. Type: Actinostachys digitata (L.) Wall. ( Acrostichum digitatum L.). Circumscription sensu Mickel & Smith (2004). Monophyletic (Wikstr€ om et al., 2002). About 15 species.
m. Acad. Roy. Sci. (Turin) 5: 419, Schizaea Sm., Me pl.9, f.9. 1793. Type: Schizaea dichotoma (L.) J.Sm. ( Acrostichum dichotomum L.). Circumscription sensu Mickel & Smith (2004). Monophyletic €m et al., 2002). Perhaps 20 species. (Wikstro 15. Family Anemiaceae Link, Fil. Spec.: 23. 1841. Circumscription sensu Smith et al. (2006b). Monophyletic (Labiak et al., 2015a). One genus and an estimated 115 species. Anemia Sw., Syn. Fil.: 6, 155. 1806. Lectotype (designated by Underw., Mem. Torrey Bot. Club 6: 267, 276. 1899): Anemia phyllitidis (L.) Sw. ( Osmunda phyllitidis L.). Circumscription sensu Mickel (2016). Includes: Mohria Sw.; Colina Greene. Monophyletic (Labiak et al., 2015a). About 115 species. L. Order Salviniales Link, Hort. Berol. 3: 155. 1833. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 1995; Pryer, 1999; Schuettpelz & Pryer, 2007; Rothfels et al., 2015). Two families, five genera, and an estimated 82 species. 16. Family Salviniaceae Martinov, Tekhno-Bot.Slovar.: 559. 1820. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer et al., 1995; Pryer, 1999; Schuettpelz & Pryer, 2007; Nagalingum et al., 2008). Two genera and an estimated 21 species. Azolla Lam., Encycl. 1(1): 343. 1783. Type: Azolla filiculoides Lam. Circumscription sensu Schneller (1990a). Monophyletic (Metzgar et al., 2007; Reid et al., 2006; Nagalingum et al., 2008). About nine species.
g., Pl. Veron. 3: 52. 1754. Type: Salvinia Salvinia Se natans (L.) All. ( Marsilea natans L.). Circumscription sensu Schneller (1990b). Monophyletic (Nagalingum et al., 2008). About 12 species. 17. Family Marsileaceae Mirb., Hist. Nat. Veg. 5: 126. 1802. Circumscription sensu Smith et al. (2006b). Monophyletic (Pryer, 1999; Schuettpelz & Pryer, 2007; Nagalingum et al., 2008). Three genera and an estimated 61 species. Marsilea L., Sp. Pl. 2: 1099. 1753. Lectotype (designated by Maxon, Sci. Surv. Porto Rico & Virgin Islands 6(3): 509. 1926): Marsilea quadrifolia L. Circumscription sensu Kramer (1990k). www.jse.ac.cn PPG I Monophyletic (Nagalingum et al., 2007, 2008; Whitten et al., 2012). About 55 species. Pilularia L., Sp. Pl. 2: 1100. 1753. Type: Pilularia globulifera L. Circumscription sensu Kramer (1990k). Monophyletic (Nagalingum et al., 2008). About five species. Regnellidium Lindm., Ark. Bot. 3(6): 2. 1904. Type: Regnellidium diphyllum Lindm. Circumscription sensu Kramer (1990k). Monotypic. M. Order Cyatheales A.B.Frank in Leunis, Syn. Pflanzenk. (ed. 2) 3: 1452. 1877. Circumscription sensu Smith et al. (2006b). Monophyletic (Korall et al., 2006; Schuettpelz et al., 2007; Rothfels et al., 2015). Eight families, 13 genera, and an estimated 713 species. 18. Family Thyrsopteridaceae C.Presl, Gef€assb€ undel Farrn: 22, 38. 1847. Circumscription sensu Smith et al. (2006b). Family consists of a single monotypic genus. Thyrsopteris Kunze, Linnaea 9: 507. 1835. Type: Thyrsopteris elegans Kunze. Circumscription sensu Kramer (1990e). Includes: Panicularia Colla. Monotypic (Korall et al., 2006). 19. Family Loxsomataceae C.Presl, Gef€assb€ undel Farrn: 31. 1847. Circumscription sensu Smith et al. (2006b). Monophyletic (Korall et al., 2006). Two monotypic genera. Loxsoma R.Br. ex Hook. in A.Cunn., Companion Bot. Mag. 2: 366. 1837. Type: Loxsoma cunninghamii R.Br. ex Hook. Circumscription sensu Kramer (1990j). Includes: Chrysodium F
ee. Monotypic. Loxsomopsis Christ, Bull. Herb. Boissier, ser. 2, 4(5): 399. 1904. Type: Loxsomopsis costaricensis Christ. Circumscription sensu Lehnert et al. (2001). Monotypic. 20. Family Culcitaceae Pic.Serm., Webbia 24: 702. 1970. Circumscription sensu Smith et al. (2006b). Monophyletic (Korall et al., 2006). One genus and two species. Culcita C.Presl, Tent. Pterid.: 135, t.5, f.5. 1836. Type: Culcita macrocarpa C.Presl. Circumscription sensu White & Turner (1988). Monophyletic (Korall et al., 2006). Two species. 21. Family Plagiogyriaceae Bower, Ann. Bot. (Oxford) 40: 484. 1926. Circumscription sensu Smith et al. (2006b). Family consists of a single genus and an estimated 15 species. Plagiogyria (Kunze) Mett., Abh. Senckenberg. Naturf. Ges. 2: 1, 268. 1858. Lectotype (designated by C.Chr., Index Filic. xliii. 1906): Plagiogyria euphlebia (Kunze) Mett. ( Lomaria euphlebia Kunze). Circumscription sensu Kramer (1990p). www.jse.ac.cn 575 Presumably monophyletic; seven species sampled to date (Korall et al., 2006; Ebihara, 2011). 15 species. 22. Family Cibotiaceae Korall, Taxon 55(3): 712. 2006. Circumscription sensu Smith et al. (2006b). Family consists of a single genus and an estimated nine species. Cibotium Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 53. 1820. Type: Cibotium chamissoi Kaulf. Circumscription sensu Kramer (1990e). Includes: Pinonia Gaudich. Monophyletic (Geiger et al., 2013). Nine species. 23. Family Metaxyaceae Pic.Serm., Webbia 24(2): 701. 1970. Circumscription sensu Smith et al. (2006b). Family consists of a single genus and an estimated six species. Metaxya C.Presl, Tent. Pterid.: 59. 1836. Type: Metaxya rostrata (Kunth) C.Presl ( Polypodium rostratum Humb. & Bonpl. ex Willd.). Circumscription sensu C
ardenas et al. (2016). Includes: Amphidesmium J.Sm.; Amphidesmium Schott ex Kunze. Monophyletic (C
ardenas et al., 2016). Six species. 24. Family Dicksoniaceae M.R.Schomb., Reis. Br.-Guiana 2: 1047. 1849. Circumscription sensu Smith et al. (2006b). Monophyletic (Korall et al., 2006; Adjie & Lestari, 2014; Noben et al., pers. comm.). Three genera and an estimated 35 species. Calochlaena (Maxon) M.D.Turner & R.A.White, Amer. Fern J. 78(3): 91. 1988. Type: Culcita dubia (R.Br.) Maxon ( Davallia dubia R.Br.). Circumscription sensu White & Turner (1988). Monophyletic (Adjie & Lestari, 2014; Noben et al., pers. comm.). Five species. Dicksonia L’H
er., Sert. Angl. 30. 1789. Lectotype (designated by J.Sm., Hist. Fil.: 254. 1875): Dicksonia
r. Circumscription sensu Kramer arborescens L’He (1990e). Includes: Balantium Kaulf. Monophyletic (Noben et al., pers. comm.). 26 species. Lophosoria C.Presl, Gef€assb€ undel Farrn: 36. 1847. Lectotype (designated by J.Sm., Hist. Fil.: 251. 1875): Lophosoria pruinata (Sw.) C.Presl ( Polypodium pruinatum Sw.). Circumscription sensu Kramer (1990i). Includes: Trichosorus Liebm. Monophyletic (Noben et al., pers. comm.). About four species. 25. Family Cyatheaceae Kaulf., Wesen Farrenkr.: 119. 1827. Circumscription sensu Smith et al. (2006b). Monophyletic (Korall et al., 2006, 2007). Three genera and an estimated 643 species. Alsophila R.Br., Prodr.: 158. 1810. Type: Alsophila australis R.Br. Circumscription sensu Conant J. Syst. Evol. 54 (6): 563–603, 2016 576 The Pteridophyte Phylogeny Group (1983). Includes: Amphicosmia Gardner; Dichorexia C.Presl; Gymnosphaera Blume; Nephelea R.M. Tryon; Thysanobotrya Alderw. Monophyletic (Korall et al., 2007). 275 species. Cyathea Sm., M
em. Acad. Roy. Sci. (Turin) 5: 416. 1793. Lectotype (designated by J.Sm., Hist. Fil.: 244. 1875): Cyathea arborea (L.) Sm. ( Polypodium arboreum L.). Circumscription sensu Korall et al. (2007). Includes: Actinophlebia C.Presl; Chnoophora Kaulf.; Cnemidaria C.Presl; Cnemidopteris Reichenb.; Cormophyllum Newm.; Disphenia C. Presl; Hemitelia R.Br.; Hemistegia C.Presl; Hymenophyllopsis K.I.Goebel; Microstegnus C.Presl; Trichopteris C.Presl; Cyathidaria Caluff. Monophyletic (Korall et al., 2007). 265 species. Sphaeropteris Bernh., J. Bot. (Schrader) 1800(2): 122. 1801. Type: Sphaeropteris medullaris (G.Forst.) Bernh. ( Polypodium medullare G.Forst.). Circumscription sensu Tryon (1970). Includes: Eatoniopteris Bomm.; Fourniera Bomm.; Schizocaena J.Sm. Monophyletic (Korall et al., 2007). 103 species. N. Order Polypodiales Link, Hort. Berol. 2: 5. 1833. Circumscription sensu Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007). Six suborders, 26 families, 253 genera, and an estimated 8714 species. Suborder Saccolomatineae Hovenkamp, this classification (see below). Circumscription equivalent to family Saccolomataceae in this classification and in Smith et al. (2006b). Additional investigation is needed to establish monophyly. One family, one genus, and an estimated 18 species. 26. Family Saccolomataceae Doweld, Phytologia 90(3): 417. 2008. Circumscription sensu Smith et al. (2006b). Contains a single genus, but additional investigation is needed (see below). An estimated 18 species. Saccoloma Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 51. 1820. Type: Saccoloma elegans Kaulf. Circumscription sensu Kramer (1990d). Includes: Orthiopteris Copel. Additional investigation is needed (Perrie et al., 2015), as indications that the genus is not monophyletic may be based on errors in sequencing or mixed samples. 18 species. Suborder Lindsaeineae Lehtonen & Tuomisto, this classification (see below). Circumscription equivalent to Lindsaeaceae in Smith et al. (2006b). Probably monophyletic, but needs to be confirmed with additional data (Lehtonen et al., 2012). Three families, nine genera, and an estimated 237 species. 27. Family Cystodiaceae J.R.Croft, Kew Bull. 41(4): 797. 1986. Circumscription sensu Christenhusz et al. (2011). Family consists of a single monotypic genus. Cystodium J.Sm, Gen. Fil. (Hooker) t.96. 1841. Type: Cystodium sorbifolium (Sm.) J.Sm. ( Dicksonia J. Syst. Evol. 54 (6): 563–603, 2016 sorbifolia Sm.). Circumscription sensu Kramer (1990e). Monotypic. 28. Family Lonchitidaceae Doweld, New Syllab. Pl. Fam.: 353. 2006. Circumscription sensu Christenhusz et al. (2011). Family consists of a single genus and an estimated two species. Lonchitis L., Sp. Pl. 2: 1078. 1753. Lectotype (designated by Brongn., Dict. Class. Hist. Nat. 9: 490. 1826): Lonchitis hirsuta L. Circumscription sensu Kramer (1990d). Presumably monophyletic, but only one species sampled to date. An estimated two species. 29. Family Lindsaeaceae C.Presl ex M.R.Schomb., Reis. Br.-Guiana 3: 883. 1849. Circumscription sensu Christenhusz et al. (2011). Monophyletic (Lehtonen et al., 2010). Seven genera and an estimated 234 species.
m. Acad. Roy. Sci. Lindsaea Dryand. ex Sm., Me (Turin) 5: 413, pl.9, f.4. 1793. Lectotype (designated by Maxon, Sci. Surv. Porto Rico & Virgin Islands 6 (3): 488. 1926): Lindsaea guianensis (Aubl.) Dryand. ( Adiantum guianense Aubl.). Circumscription sensu Lehtonen et al. (2010). Includes: Guerinia J.
e; OdonSm.; Humblotiella Tardieu; Lindsaenium Fe toloma Maxon; Sambirania Tardieu; Schizolegnia Alston; Schizoloma Gaudich.; Synaphlebium J.Sm. Monophyletic (Lehtonen et al., 2010). About 180 species. Nesolindsaea Lehtonen & Christenh., Bot. J. Linn. Soc. 163(3): 336–337. 2010. Type: Nesolindsaea caudata (Hook.) Lehtonen & Christenh. ( Lindsaea caudata Hook.). Circumscription sensu Lehtonen et al. (2010). Monophyletic (Lehtonen et al., 2010). Two species. Odontosoria F
ee, M
em. Foug., 5. Gen. Filic. 325. 1852. Type: Odontosoria uncinella (Kunze) F
ee ( Davallia uncinella Kunze). Circumscription sensu Lehtonen et al. (2010). Includes: Lindsayopsis Kuhn. Monophyletic (Lehtonen et al., 2010). About 23 species. Osmolindsaea (K.U.Kramer) Lehtonen & Christenh., Bot. J. Linn. Soc. 163(3): 335. 2010. Type: Osmolindsaea odorata (Roxb.) Lehtonen & Christenh. ( Lindsaea odorata Roxb.). Circumscription sensu Lehtonen et al. (2010, 2013). Monophyletic (Lehtonen et al., 2010, 2013). About seven species. Sphenomeris Maxon, J. Wash. Acad. Sci. 3(5): 144. 1913. Type: Sphenomeris clavata (L.) Maxon ( Adiantum clavatum L.). Circumscription sensu Lehtonen et al. (2010). Only one species sampled to date, but assumed to be monophyletic. Three species. www.jse.ac.cn PPG I Tapeinidium (C.Presl) C.Chr., Index Filic.: 631. 1906. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 53. 1947): Tapeinidium pinnatum (Cav.) C.Chr. ( Davallia pinnata Cav.). Circumscription sensu Lehtonen et al. (2010). Includes: Protolindsaya Copel. Monophyletic (Lehtonen et al., 2010). 18 species. Xyropteris K.U.Kramer, Acta Bot. Neerl. 6: 599. 1958. Type: Xyropteris stortii (Alderw.) K.U.Kramer ( Schizoloma stortii Alderw.). Circumscription sensu Lehtonen et al. (2010). Monotypic. Suborder Pteridineae J.Prado & Schuettp., this classification (see below). Circumscription equivalent to family Pteridaceae in this classification and in Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007; Rothfels et al., 2015). One family, 53 genera, and an estimated 1211 species. 30. Family Pteridaceae E.D.M.Kirchn., Schul-Bot.: 109. 1831. Circumscription sensu Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007). Five subfamilies, 53 genera, and an estimated 1211 species. Subfamily Parkerioideae Burnett, Outlines Bot.: 324. 1835, as “Parkeridae”. Circumscription equivalent to “ceratopteridoids” in Schuettpelz et al. (2007). Monophyletic (Schuettpelz et al., 2007). Two genera and an estimated nine species. Acrostichum L., Sp. Pl. 2: 1067. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 146. 1875): Acrostichum aureum L. Circumscription sensu Tryon et al. (1990). Monophyletic (Schuettpelz et al., 2007). Three species. Ceratopteris Brongn., Bull. Sci. Soc. Philom. Paris 1821: 186. 1822. Lectotype (designated by Brongn., Dict. Class. Hist. Nat. 3: 350. 1823): Ceratopteris thalictroides (L.) Brongn. ( Acrostichum thalictroides L.). Circumscription sensu Tryon et al. (1990). Monophyletic (Schuettpelz et al., 2007). Perhaps six species. 577 Jessen et al. (2012). Monophyletic (Metzgar et al., 2013). About 10 species. Llavea Lag., Gen. Sp. Pl.: 33. 1816. Type: Llavea cordifolia Lag. Circumscription sensu Tryon et al. (1990). Monotypic. Subfamily Pteridoideae Link, Fil. Spec.: 48. 1841. Circumscription sensu Zhang et al. (2015). Monophyletic (Schuettpelz et al., 2007; Zhang et al., 2015). 13 genera and an estimated 400 species. Actiniopteris Link, Fil. Spec.: 79. 1841. Lectotype (designated by Pic.Serm., Webbia 17: 6–7. 1962): Actiniopteris radiata (Sw.) Link ( Asplenium radiatum Sw.). Circumscription sensu Tryon et al. (1990). Monophyletic (Schneider et al., 2013). Six species. Anogramma Link, Fil. Spec.: 137. 1841. Lectotype (designated by C.Chr., Index Filic. xxxvii. 1906): Anogramma leptophylla (L.) Link ( Polypodium leptophyllum L.). Circumscription sensu Nakazato & Gastony (2003). Polyphyletic (Schneider et al., 2013; Hennequin et al., 2014). About five species. Austrogramme E.Fourn., Ann. Sci. Nat. Bot., ser. 5, 18: 278. 1873. Lectotype (designated by C.Chr., Index Filic. xxxvii. 1906): Austrogramme marginata (Mett.) E.Fourn. ( Gymnogramma marginata Mett.). Circumscription sensu Tryon et al. (1990). Monophyletic (Cochran et al., 2014), but relatively few species sampled. Six species. Cerosora Domin, Acta Bot. Bohem. 8: 3. 1929. Type: Cerosora chrysosora (Baker) Domin ( Gymnogramma chrysosora Baker). Circumscription sensu Schneider et al. (2013). Monophyletic (Schneider et al., 2013). Four species. Cosentinia Tod., Giorn. Sci. Nat. Econ. Palermo 1: 219. 1866. Type: Cosentinia vellea (Aiton) Tod. ( Acrostichum velleum Aiton). Circumscription sensu Tryon et al. (1990). Monotypic. Subfamily Cryptogrammoideae S.Lindsay, Edinburgh. J. Bot. 66(2): 358. 2009. Circumscription equivalent to “cryptogrammoids” in Schuettpelz et al. (2007). Monophyletic (Schuettpelz et al., 2007; Metzgar et al., 2013). Three genera and an estimated 31 species. Jamesonia Hook. & Grev., Pl. Voy. Russes Monde 2: t.178. 1830. Type: Jamesonia pulchra Hook. & Grev.
n-Mora & Gonz
alez Circumscription sensu Pabo
e; Nephopteris Lellin(2015). Includes: Eriosorus Fe
n-Mora & Gonz
alez, 2015). ger. Monophyletic (Pabo About 50 species.
e, Me
m. Foug., 5. Gen. Filic.: 167. Coniogramme Fe
e ( 1852. Type: Coniogramme javanica (Blume) Fe Gymnogramma javanica Blume). Circumscription sensu Tryon et al. (1990). Monophyletic (Metzgar et al., 2013). At least 20 species. Onychium Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 45. 1820. Type: Onychium capense Kaulf. Circumscription sensu Tryon et al. (1990). Monophyletic (Schuettpelz et al., 2007). Perhaps 10 species. Cryptogramma R.Br., Narr. Journey Polar Sea: 767. 1823. Type: Cryptogramma acrostichoides R.Br. Circumscription sensu Metzgar et al. (2013) and Pityrogramma Link, Handbuch 3: 19–20. 1833. Type: Pityrogramma chrysophylla (Sw.) Link ( Acrostichum chrysophyllum Sw.). Circumscription www.jse.ac.cn J. Syst. Evol. 54 (6): 563–603, 2016 578 The Pteridophyte Phylogeny Group sensu Tryon et al. (1990). Includes: Trismeria F
ee. Presumably monophyletic, but sampling incomplete (Schneider et al., 2013; Zhang et al., 2015). About 20 species. Pteris L., Sp. Pl. 2: 1073. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 295. 1875): Pteris longifolia L. Circumscription sensu Zhang et al. (2015). Includes: Afropteris Alston; Anopteris Prantl ex
e; Platyzoma Diels; Ochropteris J.Sm.; Neurocallis Fe R.Br. Monophyletic (Chao et al., 2014; Zhang et al., 2015). Perhaps 250 species.
e, Me
m. Soc. Hist. Nat. Strasbourg Pterozonium Fe 4(1): 202. 1850. Type: Pterozonium reniforme
e ( Gymnogramma reniformis Mart.). (Mart.) Fe Circumscription sensu Tryon et al. (1990). Monophyletic (Cochran et al., 2014), but relatively few species sampled. 14 species. Syngramma J.Sm., London J. Bot. 4: 168–169. 1845. Lectotype (designated by J.Sm., Hist. Fil.: 152. 1875): Syngramma alismifolia (C.Presl) J.Sm. ( Diplazium alismifolium C.Presl). Circumscription sensu Tryon et al. (1990). Only one species sampled to date, but assumed to be monophyletic. About 15 species. Taenitis Willd. ex Schkuhr, Kl. Linn. Pfl.-Syst. 1: 20. 1804. Type: Taenitis pteroides Willd. ex Schkuhr. Circumscription sensu Tryon et al. (1990). Monophyletic (Cochran et al., 2014), but sampling insufficient. About 15 species. Tryonia Schuettp., J.Prado & A.T.Cochran, PhytoKeys 35: 35–37. 2014. Type: Tryonia myriophylla (Sw.) Schuettp., J.Prado & A.T.Cochran ( Gymnogramma myriophylla Sw.). Circumscription sensu Cochran et al. (2014). Monophyletic (Cochran et al., 2014). Four species. Subfamily Vittarioideae Link, Fil. Spec.: 116. 1841. Circumscription equivalent to “adiantoids“ in Schuettpelz et al. (2007). Monophyletic (Prado et al., 2007; Schuettpelz et al., 2007; Rothfels & Schuettpelz, 2014). 12 genera and an estimated 345 species. Adiantum L., Sp. Pl. 2: 1094. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 274. 1875): Adiantum capillus-veneris L. Circumscription sensu Tryon et al. (1990). Monophyletic (Rothfels & Schuettpelz, 2014; Pryer et al., 2016). About 225 species. Ananthacorus Underw. & Maxon, Contr. U.S. Natl. Herb. 10: 487. 1908. Type: Ananthacorus angustifolius (Sw.) Underw. & Maxon ( Pteris angustifolia Sw.). Circumscription sensu Schuettpelz et al. (2016). Monotypic (Schuettpelz et al., 2016). Antrophyopsis (Benedict) Schuettp., Taxon 65(4): 717. 2016. Type: Antrophyopsis boryana (Willd.) J. Syst. Evol. 54 (6): 563–603, 2016 Schuettp. ( Hemionitis boryana Willd.). Circumscription sensu Schuettpelz et al. (2016). Monophyletic (Schuettpelz et al., 2016). Three species. Antrophyum Kaulf., Enum. Filic.: 197, 282. 1824. Lectotype (designated by J.Sm., Hist. Fil.: 154. 1875): Antrophyum plantagineum (Cav.) Kaulf. ( Hemionitis plantaginea Cav.). Circumscription sensu Schuettpelz et al. (2016). Monophyletic (Schuettpelz et al., 2016). Perhaps 40 species. Haplopteris C.Presl, Tent. Pterid.: 141. 1836. Type: Haplopteris scolopendrina (Bory) C.Presl ( Pteris scolopendrina Bory). Circumscription sensu Schuettpelz et al. (2016). Includes: Pleurofossa Nakai ex H.Ito; Monogramma Comm. ex Schkuhr (proposal to conserve Haplopteris against Monogramma pending; Chen et al., 2016). Monophyletic (Schuettpelz et al., 2016). Perhaps 40 species. Hecistopteris J.Sm., London J. Bot. 1: 193. 1842. Type: Hecistopteris pumila (Spreng.) J.Sm. ( Gymnogramma pumila Spreng.). Circumscription sensu Schuettpelz et al. (2016). Presumably monophyletic, but only one species sampled to date (Schuettpelz et al., 2016). Three species.
m. Soc. Linn. Paris 6: 174, Polytaenium Desv., Me 218. 1827. Type: Polytaenium lanceolatum (Sw.) Desv. ( Vittaria lanceolata Sw.). Circumscription sensu Schuettpelz et al. (2016). Includes: Anetium Splitg. Monophyletic (Schuettpelz et al., 2016). About 10 species. Radiovittaria (Benedict) E.H.Crane, Syst. Bot. 22: 514–515. 1997 (1998). Type: Radiovittaria remota (F
ee) E.H.Crane ( Vittaria remota F
ee). Circumscription sensu Schuettpelz et al. (2016). Monophyletic (Schuettpelz et al., 2016). About 10 species. Rheopteris Alston, Nova Guinea, n.s. 7: 2. 1956. Type: Rheopteris cheesmaniae Alston. Circumscription sensu Schuettpelz et al. (2016). Monotypic. Scoliosorus T.Moore, Index Fil. xxix. 1857. Type: Scoliosorus ensiformis (Hook.) T.Moore ( Antrophyum ensiforme Hook.). Circumscription sensu Schuettpelz et al. (2016). Monotypic (Schuettpelz et al., 2016).
e, Me
m. Foug., 3. Hist. Vittar.: 30–31. Vaginularia Fe 1852. Type: Vaginularia trichoidea F
ee. Circumscription sensu Schuettpelz et al. (2016). Includes: Diclidopteris Brack. Monophyletic (Schuettpelz et al., 2016). Four species.
m. Acad. Roy. Sci. (Turin) 5: 413, Vittaria Sm., Me t.9, f.5. 1793. Type: Vittaria lineata (L.) Sm. ( Pteris lineata L.). Circumscription sensu Schuettpelz et al. www.jse.ac.cn PPG I (2016). Monophyletic (Schuettpelz et al., 2016). Seven species. Subfamily Cheilanthoideae Horvat, Acta Bot. Inst. Bot. Univ. Zagreb. 2: 114. 1927, as “Cheilanthinae”. Circumscription equivalent to “cheilanthoids” in Schuettpelz et al. (2007). Monophyletic (Rothfels & Schuettpelz, 2014). 23 genera and an estimated 426 species.
e, Me
m. Foug., 5. Gen. Filic.: 145. Adiantopsis Fe 1852. Lectotype (designated by C.Chr., Index Filic.
e ( Adiantum xli. 1906): Adiantopsis radiata (L.) Fe
rez radiatum L.). Circumscription sensu Link-Pe
rez & Hickey et al. (2011) with additions by Link-Pe (2011) and Schuettpelz et al. (2014). Monophyletic
rez et al., 2011; Schuettpelz et al., 2014). (Link-Pe About 35 species.
e, Me
m. Foug., 5. Gen. Filic.: 153– Aleuritopteris Fe 154. 1852. Lectotype (designated by C.Chr., Index
e Filic. xlii. 1906): Aleuritopteris farinosa (Forssk.) Fe ( Pteris farinosa Forssk.). Circumscription sensu Zhang et al. (2013a). Includes: Leptolepidium K.H. Shing & S.K.Wu; Sinopteris C.Chr. & Ching. Not monophyletic as circumscribed (see Eiserhardt et al., 2011). About 40 species. Allosorus Bernh., Neues J. Bot. 1(2): 36. 1805. Lectotype (designated by Pic.Serm., Webbia 9: 394. 1953): Allosorus pusillus (Willd. ex Bernh.) Bernh. ( Adiantum pusillum Willd. ex Bernh.). Circumscription sensu Christenhusz (2012). Includes: Oeosporangium De Visiani. Questionably monophyletic as circumscribed. Eight species. Argyrochosma (J.Sm.) Windham, Amer. Fern J. 77 (2): 38. 1987. Lectotype (designated by C.Chr., Index Filic. xl. 1906): Argyrochosma nivea (Poir.) Windham ( Pteris nivea Poir.). Circumscription sensu Windham (1987) with additions by Wang et al. (2015). Monophyletic (Sigel et al., 2011). About 20 species. Aspidotis (Nutt. ex Hook.) Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 68. 1947. Type: Aspidotis californica (Hook.) Nutt. ex Copel. ( Hypolepis californica Hook.). Circumscription sensu Li et al. (2012b). Monophyletic (Li et al., 2012b). Four species. Astrolepis D.M.Benham & Windham, Amer. Fern J. 82(2): 55. 1992. Type: Astrolepis sinuata (Lag. ex Sw.) D.M.Benham & Windham ( Acrostichum sinuatum Lag. ex Sw.). Circumscription sensu Benham & Windham (1992) with additions by Beck et al. (2010). Monophyletic (Beck et al., 2010). Eight species. Bommeria E.Fourn., Dict. Bot. 1: 448. 1876 (1877). Lectotype (designated by Maxon, Contr. U.S. Natl. www.jse.ac.cn 579 Herb. 13: 160. 1913): Bommeria ehrenbergiana (Klotzsch) Underw. ( Gymnogramma ehrenbergiana Klotzsch). Circumscription sensu Ranker & Haufler (1990). Monophyletic based on limited sampling (two species in Eiserhardt et al., 2011). Five species. Calciphilopteris Yesilyurt & H.Schneid., Phytotaxa 7: 52. 2010. Type: Calciphilopteris ludens (Wall. ex Hook.) Yesilyurt & H.Schneid. ( Pteris ludens Wall. ex Hook.). Circumscription sensu Yesilyurt & Schneider (2010). Presumably monophyletic based on morphological similarity (only one species sampled; Schuettpelz et al., 2007; Eiserhardt et al., 2011; Rothfels & Schuettpelz, 2014). Four species. Cheilanthes Sw., Syn. Fil. 5, 126. 1806. Lectotype (designated by Maxon, Sci. Surv. Porto Rico & Virgin Islands 6(3): 428. 1926): Cheilanthes micropteris Sw. Circumscription sensu Zhang & Yatskievych (2013) but excluding Allosorus and Mildella. Includes: Cheilosoria Trevis.; Negripteris Pic.Serm.; Neurosoria Mett. ex Kuhn. Not monophyletic as circumscribed (Eiserhardt et al., 2011). About 100 species.
e, Me
m. Foug., 7. Ic. Esp. Nouv.: Cheiloplecton Fe 33, 135, t.20. 1857. Type: Cheiloplecton rigidum (Sw.) F
ee ( Pteris rigida Sw.). Circumscription sensu Mickel & Smith (2004). Monotypic (Rothfels et al., 2008; Johnson et al., 2012). Doryopteris J.Sm. emend. J.C.Yesilyurt, J. Bot. (Hooker) 3: 404. 1841. Lectotype (designated by C.V.Morton, Amer. Fern J. 34: 26. 1944): Doryopteris palmata (Willd.) J.Sm. ( Pteris palmata Willd.). Circumscription sensu Yesilyurt et al. (2015). Includes: Bakeriopteris C.Chr.; Cassebeera Kaulf.; Heteropteris F
ee; Tryonella Pic.Serm. Monophyletic (Yesilyurt et al., 2015). 21 species. Gaga Pryer, F.W.Li & Windham, Syst. Bot. 37(4): 855. 2012. Type: Gaga marginata (Kunth) F.W.Li & Windham ( Cheilanthes marginata Kunth). Circumscription sensu Li et al. (2012b). Monophyletic (Li et al., 2012b). 19 species. Hemionitis L., Sp. Pl. 2: 1077 1753. Lectotype (designated by Kaulf., Enum. Filic.: 68, 198. 1824): Hemionitis palmata L. Circumscription sensu Tryon et al. (1990). Includes: Gymnopteris Bernh.; Gymnogramma Desv.; Neurogramma Link. Monophyletic (four species sampled in Eiserhardt et al., 2011). Five species. Lytoneuron (Klotzsch) Yesilyurt, Phytotaxa 221(2): 116. 2015. Type: Lytoneuron lomariaceum (Klotzsch) Yesilyurt ( Doryopteris lomariacea Klotzsch). Circumscription sensu Yesilyurt et al. (2015). Monophyletic (Yesilyurt et al., 2015). 12 species. J. Syst. Evol. 54 (6): 563–603, 2016 580 The Pteridophyte Phylogeny Group Mildella Trevis., Reach Ist. Lombardo Sci., Rendiconti 9: 810. 1877. Type: Mildella intramarginalis (Kaulf. ex Link) Trevis. ( Pteris intramarginalis Kaulf. ex Link). Circumscription sensu Hall & Lellinger (1967). Not monophyletic as circumscribed; New World species (including type) and Old World species belong to different clades. Seven species.
e emend. Grusz & Windham, Me
m. Myriopteris Fe Foug., 5. Gen. Filic.: 148. 1852. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 65. 1947): Myriopteris marsupianthes
e. Circumscription sensu Grusz & Windham Fe (2013). Includes: Pomatophytum M.E.Jones. Monophyletic (Grusz et al., 2014). About 45 species. Notholaena R.Br., Prodr.: 145. 1810. Lectotype (designated by J.Sm., Hist. Fil.: 278. 1875): Notholaena trichomanoides (L.) Desv. ( Pteris trichomanoides L.). Circumscription sensu Rothfels et al. (2008) with additions by Yatskievych & Arbelaez (2008). Includes: Chrysochosma (J.Sm.) K€ ummerle. See Yatskievych & Smith (2003) for details regarding arguments over the typification of Notholaena. Not monophyletic as circumscribed (Rothfels et al., 2008; Johnson et al., 2012). About 30 species. Ormopteris J.Sm. ex J.Sm., Hist. Fil.: 281. 1875. Type: Ormopteris gleichenioides (Gardner) J.Sm. ( Cassebeera gleichenioides Gardner). Circumscription sensu Yesilyurt et al. (2015). Monophyletic (Yesilyurt et al., 2015). Six species. Paragymnopteris K.H.Shing, Indian Fern J. 10: 227. 1993. Type: Paragymnopteris marantae (L.) K.H. Shing ( Acrostichum marantae L.). Circumscription sensu Zhang & Ranker (2013b). Not monophyletic as circumscribed. Five species. Parahemionitis Panigrahi, Amer. Fern J. 83(3): 90, f.1. 1993. Type: Parahemionitis arifolia (Burm.f.) Panigrahi ( Asplenium arifolium Burm.f.). Circumscription sensu Zhang & Ranker (2013a). Monotypic. Pellaea Link, Fil. Spec.: 59. 1841. Lectotype (designated by C.Chr., Index Filic. xxxix. 1906): Pellaea atropurpurea (L.) Link ( Pteris atropurpurea L.). Circumscription sensu Tryon et al. (1990) but excluding Ormopteris. Includes: Choristosoria Mett. ex Kuhn; Paraceterach Copel.; Pellaeopsis J. Sm.; Platyloma J.Sm.; Pteridella Kuhn. Not monophyletic as circumscribed (Yesilyurt et al., 2015). About 40 species. Pentagramma Yatsk., Windham & E.Wollenw., Amer. Fern J. 80(1): 13. 1990. Type: Pentagramma triangularis (Kaulf.) Yatsk., Windham & E. Wollenw. ( Gymnogramma triangularis Kaulf.). J. Syst. Evol. 54 (6): 563–603, 2016 Circumscription sensu Schuettpelz et al. (2015). Monophyletic (Schuettpelz et al., 2007). Six species. Trachypteris Andr
e ex Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesamten Naturwiss. 36: 150. 1899. Type: Trachypteris aureonitens (Hook.) Andr
e ex Christ ( Acrostichum aureonitens Hook.). Circumscription sensu Ramos-Giacosa et al. (2011). Includes: Saffordia Maxon. Monophyly not assessed (only one species sampled; Eiserhardt et al., 2011). Four species. Suborder Dennstaedtiineae Schwartsb. & Hovenkamp, this classification (see below). Circumscription equivalent to family Dennstaedtiaceae in this classification and in Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007). An estimated 265 species. 31. Family Dennstaedtiaceae Lotsy, Vort. Bot. Stammesgesch. 2: 655. 1909. Circumscription sensu Smith et al. (2006b). Includes: Hypolepidaceae Pic.Serm.; Monachosoraceae Ching; Pteridiaceae Ching. Monophyletic (Schuettpelz & Pryer, 2007). Ten genera and an estimated 265 species. Blotiella R.M.Tryon, Contr. Gray Herb. 191: 96. 1962. Type: Blotiella glabra (Bory) R.M.Tryon ( Lonchitis glabra Bory). Circumscription sensu Tryon (1962) and Kramer (1990d). Monophyletic (Perrie et al., 2015), although only two species were sampled. Perhaps 20 species. Dennstaedtia Bernh., J. Bot. (Schrader) 1800(2): 124. 1801. Type: Dennstaedtia flaccida (G.Forst.) Bernh. ( Trichomanes flaccidum G.Forst.). Circumscription sensu Kramer (1990d). Includes: Adectum Link; Coptodipteris Nakai & Momose; Costaricia Christ; Emodiopteris Ching & S.K.Wu; Fuziifilix Kanai & Momose; Paradennstaedtia Tagawa; Patania C. Presl; Sitobolium Desv. Not monophyletic as circumscribed; needs further sampling to include several competing type species (Perrie et al., 2015). About 70 species. Histiopteris (J.Agardh) J.Sm., Hist. Fil.: 294: 1875. Type: Histiopteris vespertilionis (Labill.) J.Sm. ( Pteris vespertilionis Labill.). Circumscription sensu Kramer (1990d). Includes: Lepidocaulon Copel. Monophyletic (Perrie et al., 2015), although only two species were sampled. About seven species. Hypolepis Bernh., Neues J. Bot. 1(2): 34. 1805. Type: Hypolepis tenuifolia (G.Forst.) Bernh. ex C.Presl ( Lonchitis tenuifolia G.Forst.). Circumscription sensu Kramer (1990d) and Schwartsburd & Prado (2015, 2016). Mostly monophyletic (Wolf, 1995; Perrie et al., 2015), but a few species may need to be transferred to a different genus (Brownsey, 1983). Perhaps 80 species. www.jse.ac.cn PPG I Leptolepia Mett. ex Prantl, Arbeiten K€ onigl. Bot. Gart. Breslau 1: 23. 1892. Lectotype (designated by C.Chr., Index Filic. xxviii. 1906): Leptolepia novaezelandiae (Colenso) Mett. ex Diels ( Davallia novae-zelandiae Colenso). Circumscription sensu Kramer (1990d). Monotypic. Microlepia C.Presl, Tent. Pterid.: 124, pl. 4, f. 21–23. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 260. 1875): Microlepia polypodioides (Sw.) C.Presl ( Dicksonia polypodioides Sw.). Circumscription sensu Kramer (1990d). Includes: Scypholepia J.Sm. Monophyletic; sister to the old world clade of Dennstaedtia (Perrie et al., 2015). About 60 species. Monachosorum Kunze, Bot. Zeitung (Berlin) 6: 119. 1848. Type: Monachosorum davallioides Kunze. Circumscription sensu Kramer (1990m). Includes: Ptilopteris Hance; Monachosorella Hayata. Monophyletic (Perrie et al., 2015; Ebihara et al., 2016). About six species. Oenotrichia Copel., Univ. Calif. Publ. Bot. 16: 82. 1929. Type: Oenotrichia maxima (E.Fourn.) Copel. ( Leucostegia maxima E.Fourn.). Circumscription sensu Kramer (1990d). Monophyletic (Perrie et al., 2015). About two species. Paesia A.St.-Hil., Voy. Distr. Diam. 1: 381. 1833. Type: Paesia viscosa A.St.-Hil. Circumscription sensu Kramer (1990d). Monophyletic (Perrie et al., 2015), although only two species were sampled. About 15 species. Pteridium Gled. ex Scop., Fl. Carniol.: 169. 1760. Type: Pteridium aquilinum (L.) Kuhn ( Pteris aquilina L.). Circumscription sensu Kramer (1990d), Der et al. (2009), and Thomson (2012). Includes: Cincinalis Gled.; Eupteris Newman; Ornithopteris (J. Agardh) J.Sm.; Filix-foemina Farwell. Monophyletic (Der et al., 2009; Zhou et al., 2014; Wolf et al., 2015). Species number uncertain; about 20 morphotypes have been recognized, but molecular evidence points to substantially fewer species. Four species. 581 sensu Rothfels et al. (2012b). Monophyletic (Rothfels et al., 2012a, 2013). At least three species. Cystopteris Bernh., Neues J. Bot. 1(2): 26. 1805. Lectotype (designated by J.Sm., Hist. Fil.: 236. 1875): Cystopteris fragilis (L.) Bernh. ( Polypodium fragile L.). Circumscription sensu Rothfels et al. (2013) and Wei & Zhang (2014). Includes: Cystoathyrium Ching; Rhizomatopteris A.P.Khokhr. Monophyletic (Rothfels et al., 2012a, 2013; Wei & Zhang, 2014). At least 26 species. Gymnocarpium Newman, Phytologist 4: 371. 1851. Lectotype (designated by Ching, Contr. Biol. Lab. Sci. Soc. China, Bot. Ser. 9: 38. 1933): Gymnocarpium dryopteris (L.) Newman ( Polypodium dryopteris L.). Circumscription sensu Sarvela (1978) and Rothfels et al. (2013). Includes: Currania Copel. Monophyletic (Rothfels et al., 2012a, 2013). At least eight species. 33. Family Rhachidosoraceae X.C.Zhang, Phytotaxa 19: 16. 2011. Circumscription sensu Christenhusz et al. (2011) and Rothfels et al. (2012b). Family consists of a single genus and an estimated eight species. Rhachidosorus Ching, Acta Phytotax. Sin. 9(1): 73. 1964. Type: Rhachidorous mesosorus (Makino) Ching ( Asplenium mesosorum Makino). Circumscription sensu Lindsay et al. (2012). Likely monophyletic (Li et al., 2011; Rothfels et al., 2012a). Eight species. 34. Family Diplaziopsidaceae X.C.Zhang & Christenh., Phytotaxa 19: 15. 2011. Circumscription sensu Rothfels et al. (2012b). Likely monophyletic (Kuo et al., 2011; Rothfels et al., 2012a). Two genera and an estimated four species. Diplaziopsis C.Chr., Index Filic. 227. 1905. Type: Diplaziopsis brunoniana (Wall.) W.M.Chu ( Allantodia brunoniana Wall.). Circumscription sensu He & Kato (2013). Likely monophyletic (Li et al., 2011; Wei et al., 2010; Rothfels et al., 2012a). Three species. Suborder Aspleniineae H.Schneid. & C.J.Rothf., this classification (see below). Circumscription equivalent to “eupolypods II” in Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007; Rothfels et al., 2012a). 11 families, 72 genera, and an estimated 2775 species. Homalosorus Small ex Pic.Serm., Webbia 31(1): 246. 1977. Type: Homalosorus pynocarpos (Spreng.) Pic. Serm. ( Asplenium pycnocarpum Spreng.). Circumscription sensu Pichi Sermolli (1977a). Monotypic. 32. Family Cystopteridaceae Shmakov, Turczaninowia 4: 60. 2001. Circumscription sensu Rothfels et al. (2012b). Monophyletic (Rothfels et al., 2012a, 2013; Wei & Zhang, 2014). Three genera and an estimated 37 species. 35. Family Desmophlebiaceae Mynssen, A.Vasco, Sylvestre, R.C.Moran & Rouhan, Taxon 65(1): 19. 2016. Circumscription sensu Mynssen et al. (2016). Family consists of a single genus and two species. Acystopteris Nakai, Bot. Mag. (Tokyo) 47: 180. 1933. Type: Acystopteris japonica (Luerss.) Nakai ( Cystopteris japonica Luerss.). Circumscription www.jse.ac.cn Desmophlebium Mynssen, A.Vasco, Sylvestre, R.C. Moran & Rouhan, Taxon 65(1): 27, f.5. 2016. Type: Desmophlebium lechleri (Mett.) Mynssen, A.Vasco, Sylvestre, R.C.Moran & Rouhan ( Asplenium J. Syst. Evol. 54 (6): 563–603, 2016 582 The Pteridophyte Phylogeny Group lechleri Mett.). Circumscription sensu Mynssen et al. (2016). Only one species included in a phylogenetic analysis, but assumed to be monophyletic based on morphological evidence (Mynssen et al., 2016). Two species. 36. Family Hemidictyaceae Christenh. & H.Schneid., Phytotaxa 28: 51. 2011. Circumscription sensu Rothfels et al. (2012b) and Christenhusz & Schneider (2011). Family consists of a single monotypic genus. Hemidictyum C.Presl, Tent. Pterid.: 110, t.3, f.24–26. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 335. 1875): Hemidictyum marginatum (L.) C.Presl ( Asplenium marginatum L.). Circumscription sensu Kramer et al. (1990). Monotypic. 37. Family Aspleniaceae Newman, Hist. Brit. Ferns: 6. 1840. Circumscription sensu Smith et al. (2006b) and Rothfels et al. (2012b). Monophyletic (Schneider et al., 2004a; Schneider et al., 2005). Two genera and an estimated 730 species. Asplenium L., Sp. Pl. 2: 1078. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 316. 1875): Asplenium marinum L. Circumscription sensu Schneider et al. (2004a). Includes: Antigramma C.Presl; Camptosorus Link; Ceterach Willd.; Ceterachopsis (J.Sm.) Ching; Diellia Brack.; Diplora Baker; Holodictyum Maxon; Loxoscaphe T.Moore; Phyllitis Hill; Pleurosorus F
ee; Schaffneria F
ee ex T.Moore; Scolopendrium Adans.; Sinephropteris Mickel. Monophyletic (Schneider et al., 2004a). At least 700 species. Hymenasplenium Hayata, Bot. Mag. (Tokyo) 41 (492): 712. 1927. Type: Hymenasplenium unilaterale (Lam.) Hayata ( Asplenium unilaterale Lam.). Circumscription sensu Schneider et al. (2004a). Monophyletic (Schneider et al., 2004a). At least 30 species. 38. Family Woodsiaceae Herter, Revista Sudamer. Bot. 9: 14. 1949. Circumscription sensu Christenhusz et al. (2011) and Rothfels et al. (2012b). Family consists of a single genus and an estimated 39 species. Woodsia R.Br., Prodr.: 158, Obs. 4. 1810. Lectotype (designated by J.Sm., Hist. Fil.: 237. 1875): Woodsia ilvensis (L.) R.Br. ( Acrostichum ilvense L.). Circumscription sensu Shao et al. (2015). Includes: Eriosoriopsis (Kitag.) Ching & S.H.Wu; Cheilanthopsis Hieron.; Hymenocystis C.A.Mey; Physematium Kaulf.; Protowoodsia Ching; Woodsiopsis Shmakov. Monophyletic (Rothfels et al., 2012a; Shao et al., 2015; Shmakov, 2015). 39 species. 39. Family Onocleaceae Pic.Serm., Webbia 24: 708. 1970. Circumscription sensu Smith et al. (2006b). Monophyletic (Gastony & Ungerer, 1997; Rothfels et al., 2012a). Four genera and an estimated five species. J. Syst. Evol. 54 (6): 563–603, 2016 Matteuccia Tod., Giorn. Sci. Nat. Econ. Palermo 1 (3–4): 235. 1866. Type: Matteuccia struthiopteris (L.) Tod. ( Osmunda struthiopteris L.). Circumscription sensu Gastony & Ungerer (1997) and Rothfels et al. (2012b). Monotypic, although American and European populations could be treated as distinct species (Koenemann et al., 2011). Onoclea L., Sp. Pl. 2: 1062. 1753. Type: Onoclea sensibilis L. Circumscription sensu Gastony & Ungerer (1997) and Rothfels et al. (2012b). Includes: Pterinodes Siegesb. ex Kuntze; Calypterium Bernh.; Ragiopteris C.Presl; Riedlea Mirb. Monotypic (Gastony & Ungerer, 1997). Onocleopsis F.Ballard, Amer. Fern J. 35: 1. 1945. Type: Onocleopsis hintonii F.Ballard. Circumscription sensu Gastony & Ungerer (1997) and Rothfels et al. (2012b). Monotypic. Pentarhizidium Hayata, Bot. Mag. (Tokyo) 42: 345. 1928. Type: Pentarhizidium japonicum Hayata. Circumscription sensu Gastony & Ungerer (1997) and Rothfels et al. (2012b). Monophyletic (Gastony & Ungerer, 1997; Rothfels et al., 2012a). The name Pentarhizidium was first used in 1927, in the context of describing the morphological distinctiveness of the P. orientale stele (Hayata, 1927). However, the paper lacked a description of the genus itself and we consider Pentarhizidium and P. orientale as nomina nuda until the genus was formally described and a type (P. japonicum) was designated (Hayata, 1928). Two species. 40. Family Blechnaceae Newman, Hist. Brit. Ferns (ed. 2): 8. 1844. Circumscription sensu Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer 2007; Rothfels et al., 2012a; Gasper et al., 2016a). Three subfamilies, 24 genera, and an estimated 265 species. Subfamily Stenochlaenoideae (Ching) J.P.Roux, Conspect. South. Afr. Pteridophyta: 156. 2001. Circumscription sensu Roux (2001) emend. Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). Three genera and an estimated 12 species. Salpichlaena J.Sm., Gen. Fil. pl. 93. 1842. Type: Salpichlaena volubilis (Kaulf.) J.Sm. ( Blechnum volubile Kaulf.). Circumscription sensu Perrie et al. (2014) and Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). Three species. Stenochlaena J.Sm., J. Bot. (Hooker) 3: 401. 1841. Lectotype (designated by Pfeiffer, Nom. 2: 1274. 1874): Stenochlaena palustris (Burm.f.) Bedd. ( Polypodium palustre Burm.f.). Circumscription sensu Perrie et al. (2014) and Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). We follow Copeland (1947) in identifying the name Acrostichum scandens L. (referred to in both the www.jse.ac.cn PPG I 583 description of the genus and its lectotypification) with Stenochlaena palustris (Burm.f.) Bedd. Seven species. Blechnum orientale L.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). Two species. Telmatoblechnum Perrie, D.J.Ohlsen & Brownsey, Taxon 63(4). 755. 2014. Type: Telmatoblechnum serrulatum (Rich.) Perrie, D.J.Ohlsen & Brownsey ( Blechnum serrulatum Rich.). Circumscription sensu Perrie et al. (2014) and Gasper et al. (2016b). Monophyletic (Perrie et al., 2014; Gasper et al., 2016a). Two species. Blechnum L., Sp. Pl. 2: 1077. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 300. 1875): Blechnum occidentale L. Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 30 species. Subfamily Woodwardioideae Gasper, V.A.O.Dittrich & Salino, Phytotaxa 275(3): 197. 2016. Circumscription sensu Gasper et al. (2016b). Monophyletic (Cranfill & Kato, 2003; Perrie et al., 2014; Gasper et al., 2016a). Three genera and an estimated 15 species. Anchistea C.Presl, Epimel. Bot.: 71. 1851. Type: Anchistea virginica (L.) C.Presl ( Blechnum virginicum L.). Circumscription sensu Gasper et al. (2016b). Monotypic (Gasper et al., 2016a). Lorinseria C.Presl, Epimel. Bot.: 72. 1851. Lectotype (designated by J.Sm., Hist. Fil.: 310. 1875): Lorinseria areolata (L.) C.Presl ( Acrostichum areolatum L.). Circumscription sensu Gasper et al. (2016b). Lorinseria C.Presl appears to be a later homonym of Lorinsera Opiz, and will therefore most likely require a new name (or conservation). Monotypic (Gasper et al., 2016a). Woodwardia Sm., M
em. Acad. Roy. Sci. Turin 5. 411. 1793. Lectotype (designated by J.Sm., Hist. Fil.: 310. 1875): Woodwardia radicans (L.) Sm. ( Blechnum radicans L.). Circumscription sensu Gasper et al. (2016b). Includes: Chieniopteris Ching. Monophyletic (Gasper et al., 2016a). 13 species. Subfamily Blechnoideae Gasper, V.A.O.Dittrich & Salino, Phytotaxa 275(3): 201. 2016. Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 18 genera and an estimated 238 species. Austroblechnum Gasper & V.A.O.Dittrich, Phytotaxa 275(3): 202. 2016. Type: Austroblechnum penna-marina (Poir.) Gasper & V.A.O.Dittrich ( Polypodium penna-marina Poir.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 40 species. Blechnidium T.Moore, Index Fil. clv. 1860; Brit. Ferns: 210. 1860. Type: Blechnidium melanopus (Hook.) T.Moore ( Blechnum melanopus Hook.). Circumscription sensu Gasper et al. (2016b). Monotypic (Gasper et al., 2016a). Blechnopsis C.Presl, Epimel. Bot.: 115. 1851. Lectotype (designated by Pic.Serm., Webbia 28: 456. 1973): Blechnopsis orientalis (L.) C.Presl ( www.jse.ac.cn Brainea J.Sm., Cat. Kew Ferns: 5. 1856. Type: Brainea insignis (Hook.) J.Sm. ( Bowringia insignis Hook.). Circumscription sensu Gasper et al. (2016b). Monotypic (Gasper et al., 2016a). Cleistoblechnum Gasper & Salino, Phytotaxa 275 (3): 207. 2016. Type: Cleistoblechnum eburneum (Christ) Gasper & Salino ( Blechnum eburneum Christ). 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.) Gasper & V.A.O.Dittrich ( Stegania fluviatilis R.Br.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 11 species. Diploblechnum Hayata, Bot. Mag. (Tokyo) 41. 702. 1927. Type: Diploblechnum integripinnulum Hayata. Circumscription sensu Gasper et al. (2016b). Includes: Pteridoblechnum Hennipman and Steenisioblechnum Hennipman. Monophyletic (Gasper et al., 2016a). Six species. Doodia R.Br., Prodr. Fl. Nov. Holl.: 151. 1810. Lectotype (designated by J.Sm., Hist. Fil.: 309. 1875): Doodia aspera R.Br. Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 19 species. Icarus Gasper & Salino, Phytotaxa 275(3): 209. 2016. Type: Icarus filiformis (A.Cunn.) Gasper & Salino ( Lomaria filiformis A.Cunn.). Circumscription sensu Gasper et al. (2016b). Monotypic (Gasper et al., 2016a). Lomaria Willd., Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 3: 160. 1809. Lectotype (designated by J.Sm., Hist. Fil.: 303. 1875): Lomaria nuda (Labill.) Willd. ( Onoclea nuda Labill.). Circumscription sensu Gasper et al. (2016b). Includes: Stegania R.Br. Monophyletic (Gasper et al., 2016a). Six species. Lomaridium C.Presl, Epimel. Bot.: 154. 1851. Lectotype (designated by C.Chr., Index Filic. xxxv. 1906): Lomaridium plumieri (Desv.) C.Presl ( Lomaria plumieri Desv.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 16 species. J. Syst. Evol. 54 (6): 563–603, 2016 584 The Pteridophyte Phylogeny Group Lomariocycas (J.Sm.) Gasper & A.R.Sm., Phytotaxa 275(3): 212. 2016. Type: Lomaria boryana (Sw.) Willd. ( Onoclea boryana Sw.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 19 species. Neoblechnum Gasper & V.A.O.Dittrich, Phytotaxa 275(3): 214. 2016. Type: Neoblechnum brasiliense (Desv.) Gasper & V.A.O.Dittrich ( Blechnum brasiliense Desv.). Circumscription sensu Gasper et al. (2016b). Monotypic (Gasper et al., 2016a). Oceaniopteris Gasper & Salino, Phytotaxa 275(3): 214. 2016. Type: Oceaniopteris gibba (Labill.) Gasper & Salino ( Lomaria gibba Labill.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). Eight species. Parablechnum C.Presl, Epimel. Bot.: 109. 1851.
e, Me
m. Foug., 5. Lectotype (designated by Fe Gen. Filic.: 82. 1852): Parablechnum procerum (G. Forst.) C.Presl ( Osmunda procera G.Forst.). Circumscription sensu Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). 65 species. Sadleria Kaulf., Enum. Filic.: 161. 1824. Type: Sadleria cyatheoides Kaulf. Circumscription sensu Perrie et al. (2014) and Gasper et al. (2016b). Monophyletic (Gasper et al., 2016a). Six species. Struthiopteris Scop., Fl. Carn.: 168. 1760. Lectotype (designated by Weis, Pl. Crypt. Fl. Gott.: 286. 1770): Struthiopteris spicant (L.) Weis ( Lomaria spicant L.). Circumscription sensu Gasper et al. (2016b). Includes: Spicantopsis Nakai. Monophyletic (Gasper et al., 2016a). Five species. 41. Family Athyriaceae Alston, Taxon 5: 25. 1956. Circumscription sensu Rothfels et al. (2012b). Monophyletic (Rothfels et al., 2012a). Three genera and an estimated 650 species. Athyrium Roth, Tent. Fl. Germ. 3(1): 31. 1800. Lectotype (designated by J.Sm., Hist. Fil.: 327. 1875): Athyrium filix-femina (L.) Roth ( Polypodium filix-femina L.). Circumscription equivalent to “athyriids“ in Rothfels et al. (2012b). Includes: Anisocampium C.Presl; Kuniwatsukia Pic.Serm.; Pseudocystopteris Ching; Neoathyrium Ching & Z.R.Wang; Cornoathyrium Nakaike; Cornopteris Nakai. Monophyletic (Adjie et al., 2008; Rothfels et al., 2012a). About 230 species. Deparia Hook. & Grev., Icon. Filic. 2(8): pl.154. 1830. Type: Deparia macraei Hook. & Grev. Circumscription sensu Kuo et al. (2016b). Includes: Depazium Nakaike; Lunathyrium Koidz.; Dryoathyrium Ching; Parathyrium Holttum; Athyriopsis Ching; Dictyodroma Ching; Triblemma (J.Sm.) Ching. Monophyletic (Kuo et al., 2016b). About 70 species. J. Syst. Evol. 54 (6): 563–603, 2016 Diplazium Sw., J. Bot. (Schrader) 1800(2): 4, 61. 1801. Lectotype (designated by J.Sm., Hist. Fil.: 325. 1875): Diplazium plantaginifolium (L.) Urb. ( Asplenium plantaginifolium L.). Circumscription sensu Wei et al. (2013). Includes: Allantodia R.Br.; Anisogonium C.Presl; Callipteris Bory; Monomelangium Hayata. Monophyletic (Wei et al., 2013, 2015). About 350 species. 42. Family Thelypteridaceae Ching ex Pic.Serm., Webbia 24: 709. 1970. Circumscription sensu Smith & Cranfill (2002), He & Zhang (2012), and Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; Rothfels et al., 2012a; Almeida et al., 2016). Two subfamilies, 30 genera, and an estimated 1034 species. Subfamily Phegopteridoideae Salino, A.R.Sm. & T.E.Almeida, this classification (see below). Circumscription sensu Smith & Cranfill (2002), He & Zhang (2012), and Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Rothfels et al., 2012a; Almeida et al., 2016). Three genera and an estimated 34 species. Macrothelypteris (H.Ito) Ching, Acta Phytotax. Sin. 8(4): 308. 1963. Type: Macrothelypteris oligophlebia (Baker) Ching ( Nephrodium oligophlebium Baker). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 10 species. Phegopteris (C.Presl) F
ee, M
em. Foug., 5. Gen. Filic.: 242. 1852. Lectotype (designated by Ching, Acta Phytotax. Sin. 8(4): 312. 1963): Phegopteris polypodioides F
ee ( Polypodium phegopteris L.). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). For discussion of the authority and typification, see McNeill and Pryer (1985). Four species. Pseudophegopteris Ching, Acta Phytotax. Sin. 8(4): 313. 1963. Type: Pseudophegopteris pyrrhorachis (Kunze) Ching ( Polypodium pyrrhorachis Kunze). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 20 species. Subfamily Thelypteridoideae C.F.Reed, Phytologia 17: 254. 1968. Circumscription sensu Smith & Cranfill (2002), He & Zhang (2012), and Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 27 genera and an estimated 1000 species. Amauropelta Kunze, Farrnkr€auter 1: 86. 1843. Type: Amauropelta breutelii Kunze. Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; Almeida et al., 2016). About 215 species. www.jse.ac.cn PPG I Amblovenatum J.P.Roux, Strelitzia 23: 200. 2009. Replacement name for Amphineuron Holttum non Amphineurion (A.DC.) Pichon. Type: Amblovenatum opulentum (Kaulf.) J.P.Roux ( Aspidium opulentum Kaulf.). Circumscription sensu Almeida et al. (2016). Monophyletic (He & Zhang, 2012; Almeida et al., 2016). 12 species. 585 Meniscium Schreb., Gen. Pl. ed. 8(a) 2: 757. 1791. Type: Meniscium reticulatum (L.) Sw. ( Polypodium reticulatum L.). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 27 species. Ampelopteris Kunze, Bot. Zeitung (Berlin) 6: 114. 1848. Type: Ampelopteris elegans Kunze. Circumscription sensu Almeida et al. (2016). Monotypic (He & Zhang, 2012; Almeida et al., 2016). Menisorus Alston, Bol. Soc. Brot., s
er. 2. 30: 20. 1956. Type: Menisorus pauciflorus (Hook.) Alston ( Meniscium pauciflorum Hook.). Circumscription sensu Holttum (1974a). Not yet included in a phylogenetic study. Monotypic. Chingia Holttum, Blumea 19(1): 31. 1971. Type: Chingia ferox (Blume) Holttum ( Aspidium ferox Blume). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 20 species. Mesophlebion Holttum, Blumea 19(1): 29. 1971. Type: Mesophlebion crassifolium (Blume) Holttum ( Aspidium crassifolium Blume). Circumscription sensu Holttum (1975). Monophyly uncertain (only one species sampled; Almeida et al., 2016). 15 species.
v., Fl. Kouy-Tche
ou 472. 1915. LectoChristella H.Le type (designated by Holttum, Taxon 20: 534. 1971): Christella parasitica (L.) H.Lev. ( Polypodium parasiticum L.). Circumscription sensu Almeida et al. (2016). Not monophyletic as circumscribed (Almeida et al., 2016). Perhaps 70 species. Mesopteris Ching, Acta Phytotax. Sin. 16(4): 21. 1978. Type: Mesopteris tonkinensis (C.Chr.) Ching ( Dryopteris tonkinensis C.Chr.). Circumscription sensu Lin et al. (2013). Monotypic. Coryphopteris Holttum, Blumea 19(1): 33. 1971. Type: Coryphopteris viscosa (Baker) Holttum ( Nephrodium viscosum Baker). Circumscription sensu Almeida et al. (2016). Includes: Parathelypteris Ching (pro parte). Monophyletic (Smith & Cranfill, 2002; Almeida et al., 2016). 47 species. Cyclogramma Tagawa, Acta Phytotax. Geobot. 7 (1): 52. 1938. Type: Cyclogramma simulans (Ching) Tagawa ( Thelypteris simulans Ching). Circumscription sensu Almeida et al. (2016). Presumably monophyletic (He & Zhang, 2012; Almeida et al., 2016). Eight species. Cyclosorus Link, Hort. Berol. 2: 128. 1833. Type: Cyclosorus gongylodes (Schkuhr) Link ( Aspidium gongylodes Schkuhr). Circumscription sensu Almeida et al. (2016). Monophyletic (Almeida et al., 2016). Two species. Glaphyropteridopsis Ching, Acta Phytotax. Sin. 8 (4): 320. 1963. Type: Glaphyropteridopsis erubescens (Wall. ex Hook.) Ching ( Polypodium erubescens Wall. ex Hook.). Circumscription sensu Holttum (1971). Presumably monophyletic (Almeida et al., 2016). Four species. Goniopteris C.Presl, Tent. Pterid.: 181, pl.7, f.9–11. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 191. 1875): Goniopteris crenata (Sw.) C.Presl ( Polypodium crenatum Sw.). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). About 120 species. www.jse.ac.cn Metathelypteris (H.Ito) Ching, Acta Phytotax. Sin. 8(4): 305. 1963. Type: Metathelypteris gracilescens (Blume) Ching ( Aspidium gracilescens Blume). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 12 species. Nannothelypteris Holttum, Blumea 19(1): 38. 1971. Type: Nannothelypteris aoristisora (Harr.) Holttum ( Polypodium aoristisorum Harr.). Circumscription sensu Holttum (1971). Monophyly uncertain (only one species sampled; Almeida et al., 2016). Three species. Oreopteris Holub, Folia Geobot. Phytotax. 4 (1): 46. 1969. Type: Oreopteris limbosperma (All.) Holub ( Polypodium limbospermum All.). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). Three species. Parathelypteris (H.Ito) Ching, Acta Phytotax. Sin. 8: 300. 1963. Type: Parathelypteris glanduligera (Kunze) Ching ( Aspidium glanduligerum Kunze). Circumscription sensu Holttum (1971). Not monophyletic as currently circumscribed (Almeida et al., 2016). About 15 species. Plesioneuron (Holttum) Holttum, Blumea 22: 232. 1975. Type: Plesioneuron tuberculatum (Ces.) Holttum ( Nephrodium tuberculatum Ces.). Circumscription sensu Holttum (1975). Monophyly uncertain (only one species sampled; Almeida et al., 2016). 30 species. J. Syst. Evol. 54 (6): 563–603, 2016 586 The Pteridophyte Phylogeny Group Pneumatopteris Nakai, Bot. Mag. (Tokyo) 47: 179. 1933. Type: Pneumatopteris callosa (Blume) Nakai ( Aspidium callosum Blume). Circumscription sensu Holttum (1971). Monophyly uncertain (only two species sampled; He & Zhang, 2012; Almeida et al., 2016). About 80 species. Pronephrium C.Presl, Epimel. Bot.: 258. 1851. Lectotype (designated by Holttum, Novit. Bot. 1968: 48): Pronephrium lineatum (Blume) C.Presl ( Aspidium lineatum Blume). Circumscription
e; sensu Holttum (1971). Includes: Abacopteris Fe Dimorphopteris Tagawa & K.Iwats. Not monophyletic as circumscribed (Almeida et al., 2016). 68 species. Pseudocyclosorus Ching, Acta Phytotax. Sin. 8(4): 322. 1963. Type: Pseudocyclosorus tylodes (Kunze) Ching ( Aspidium tylodes Kunze). Circumscription sensu Holttum & Grimes (1980). Not monophyletic as circumscribed (Almeida et al., 2016). 11 species. Sphaerostephanos J.Sm., Gen. Fil. (Hooker): pl.24. 1840. Type: Sphaerostephanos asplenioides J.Sm. Circumscription sensu Holttum (1971). Not monophyletic as circumscribed (Almeida et al., 2016). About 185 species. Stegnogramma Blume, Enum. Pl. Javae 2: 172. 1828. Type: Stegnogramma aspidioides Blume. Circumscription sensu Almeida et al. (2016). Includes: Dictyocline T.Moore; Leptogramma J. Sm. Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 18 species. Steiropteris (C.Chr.) Pic.Serm., Webbia 28: 449. 1973. Lectotype (designated by C.Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 1913: 164): Steiropteris deltoidea (Sw.) Pic.Serm. ( Polypodium deltoideum Sw.). Circumscription sensu Almeida et al. (2016). In
e. Monophyletic (Smith cludes: Glaphyropteris Fe & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). 22 species. Thelypteris Schmidel, Icon. Pl. 3: 45–48, pl.11, 13. 1763. Type: Thelypteris palustris Schott ( Acrostichum thelypteris L.). Circumscription sensu Almeida et al. (2016). Monophyletic (Smith & Cranfill, 2002; He & Zhang, 2012; Almeida et al., 2016). Two species. Trigonospora Holttum, Blumea 19(1): 29. 1971. Type: Trigonospora ciliata (Benth.) Holttum ( Aspidium ciliatum Benth.). Circumscription sensu Holttum & Grimes (1980). Monophyly uncertain (only one species sampled; Almeida et al., 2016). Eight species. J. Syst. Evol. 54 (6): 563–603, 2016 Suborder Polypodiineae Dumort., Anal. Fam. Pl.: 67. 1829. Circumscription equivalent to “eupolypods I“ in Smith et al. (2006b). Monophyletic (Schuettpelz & Pryer, 2007; Rothfels et al., 2015). Nine families, 108 genera, and an estimated 4208 species. 43. Family Didymochlaenaceae Ching ex Li Bing Zhang & Liang Zhang, Taxon 64(1): 34. 2015. Circumscription sensu Zhang & Zhang (2015). Family consists of a single monotypic genus. Didymochlaena Desv., Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 5: 303. 1811. Type: Didymochlaena sinuosa Desv. Circumscription sensu Zhang & Zhang (2015). Monotypic, although undescribed species may exist. 44. Family Hypodematiaceae Ching, Acta Phytotax. Sin. 13 (1): 96. 1975. Circumscription sensu Zhang & Zhang (2015). Monophyletic (Zhang & Zhang, 2015). Two genera and an estimated 22 species. Hypodematium Kunze, Flora 16(2): 690. 1833. Type: Hypodematium onustum Kunze. Circumscription sensu Kramer et al. (1990). Monophyletic (Zhang & Zhang, 2015). About 20 species. Leucostegia C.Presl, Tent. Pterid.: 94, pl.4, f.11. 1836. Type: Leucostegia immersa C.Presl. Circumscription sensu Kramer (1990c). Monophyletic (Zhang & Zhang, 2015). Two species. 45. Family Dryopteridaceae Herter, Revista Sudamer. Bot. 9(1): 15. 1949. Circumscription sensu Liu et al. (2016). Monophyletic (Liu et al., 2016). Three subfamilies, 26 genera, and an estimated 2115 species. Subfamily Polybotryoideae H.M.Liu & X.C.Zhang, Pl. Syst. Evol. 302: 330. 2016. Circumscription sensu Liu et al. (2016). Monophyletic (Moran & Labiak, 2015; Liu et al., 2016). Seven genera and an estimated 98 species. Cyclodium C.Presl, Tent. Pterid.: 85, pl.2, f.20–21. 1836. Lectotype (designated by C.Chr., Index Filic. xxv. 1906): Cyclodium meniscioides (Willd.) C.Presl ( Aspidium meniscioides Willd.). Circumscription sensu Smith (1986) and Moran & Labiak (2015). Monophyletic (Moran & Labiak, 2015). About 10 species. Maxonia C.Chr., Smithsonian Misc. Collect. 66(9): 3. 1916. Type: Maxonia apiifolia (Sw.) C.Chr. ( Dicksonia apiifolia Sw.). Circumscription sensu Christensen (1916) and Moran & Labiak (2015). Monotypic (Prado & Moran, 2016). Olfersia Raddi, Opusc. Sci. 3: 283, t.11, f.b. 1819. Type: Olfersia corcovadensis Kaulf. ex Raddi. Circumscription sensu Moran (1986) and Moran www.jse.ac.cn PPG I & Labiak (2015). Monophyletic (Moran & Labiak, 2015). Three species. Polybotrya Humb. & Bonpl. ex Willd., Sp. Pl. 5(1): 99. 1810. Type: Polybotrya osmundacea Humb. & Bonpl. ex Willd. Circumscription sensu Moran (1987) and Moran & Labiak (2015). Monophyletic (Moran & Labiak, 2015). 35 species. Polystichopsis (J.Sm.) C.Chr., Verdoorn, Man. Pterid.: 543. 1938. Lectotype (designated by C. Chr., Index Filic. Suppl. 3, 7. 1934): Polystichopsis pubescens (L.) C.V.Morton ( Polypodium pubescens L.). Circumscription sensu Moran & Labiak (2015) and Prado & Moran (2016). Monophyletic (Moran & Labiak, 2015). Eight species. Stigmatopteris C.Chr., Bot. Tidsskr. 29: 292. 1909. Type: Stigmatopteris rotundata (Willd.) C.Chr. ( Aspidium rotundatum Willd.). Circumscription sensu Moran (1991) and Moran & Labiak (2015, 2016). Monophyletic (Moran & Labiak, 2016). 40 species. Trichoneuron Ching, Acta Phytotax. Sin. 10(2): 118, pl. 22. 1965. Type: Trichoneuron microlepioides Ching. Circumscription sensu Liu et al. (2016). Monotypic (Liu et al., 2016). Subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, Jermy & Mickel, Fern Gaz. 11: 154. 1975. Circumscription sensu Zhang et al. (2013b) and Liu et al. (2016). Monophyletic (Liu et al., 2016). 11 genera and an estimated 883 species. Arthrobotrya J.Sm., Hist. Fil.: 141. 1875. Type: Arthrobotrya articulata (F
ee) J.Sm. ( Polybotrya articulata F
ee). Circumscription sensu Moran et al. (2010a). Monophyletic (Labiak et al., 2014). Two species. Bolbitis Schott, Gen. Fil.: pl.14. 1834. Lectotype (designated by C.Chr., Index Filic. xxvi. 1906): Bolbitis serratifolia (Mert. ex Kaulf.) Schott ( Acrostichum serratifolium Mert. ex Kaulf.). Circumscription sensu Moran et al. (2010a). Includes: Campium C.Presl; Cyrtogonium J.Sm.; Edanyoa
e; Copel.; Egenolfia Schott; Heteroneuron Fe Jenkinsia Hook.; Poecilopteris C.Presl. Monophyletic (Moran et al., 2010a, 2010b). About 80 species. Elaphoglossum Schott ex J.Sm., J. Bot. (Hooker) 4: 148. 1842. Lectotype (designated by J.Sm., Hist. Fil.: 125. 1875): Elaphoglossum conforme (Sw.) J.Sm. ( Acrostichum conforme Sw.). Circumscription sensu Moran et al. (2010a). Includes: Aconiopteris C.Presl;
e; MicrostaDictyoglossum Sm.; Hymenodium Fe phyla C.Presl; Peltapteris Link; Rhipidopteris Schott
e. Monophyletic (Rouhan et al., 2004; Lo
riga ex Fe et al., 2014). About 600 species. www.jse.ac.cn 587 Lastreopsis Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (4): 157. 1938. Type: Lastreopsis recedens (J.Sm. ex T.Moore) Ching ( Lastrea recedens J.Sm. ex T. Moore). Circumscription sensu Tindale (1965) and Labiak et al. (2014, 2015b). Includes: Coveniella Tindale. See also Labiak et al. (2015b). Monophyletic (Labiak et al., 2014). 16 species. Lomagramma J.Sm., J. Bot. (Hooker) 3: 402. 1841. Type: Lomagramma pteroides J.Sm. Circumscription sensu Moran et al. (2010a). Includes: Cheilo
e. Monophyletic (Moran et al., 2010a). 15 lepton Fe species. Megalastrum Holttum, Gard. Bull. Singapore 39(2): 161. 1986. Type: Megalastrum villosum (L.) Holttum ( Polypodium villosum L.). Circumscription sensu Holttum (1986) and Liu et al. (2016). Monophyletic (Labiak et al., 2014). 91 species. Mickelia R.C.Moran, Labiak & Sundue, Brittonia 62 (4): 338. 2010. Type: Mickelia nicotianifolia (Sw.) R.C.Moran, Labiak & Sundue ( Acrostichum nicotianifolium Sw.). Circumscription sensu Moran et al. (2010a, 2010b). Monophyletic (Moran et al., 2010b). 10 species. Parapolystichum (Keyserl.) Ching, Sunyatsenia 5: 239. 1940. Type: Parapolystichum effusum (Sw.) Ching ( Polypodium effusum Sw.). Circumscription sensu Labiak et al. (2014, 2015b). Monophyletic (Labiak et al., 2014). See also Labiak et al. (2015b) and Sundue & Testo (2016). 28 species. Pleocnemia C.Presl, Tent. Pterid.: 183, pl.7, f.12. 1836. Type: Pleocnemia leuzeana (Gaudich.) C.Presl ( Polypodium leuzeanum Gaudich.). Circumscription sensu Holttum (1974b) and Liu et al. (2016). Monophyletic (Labiak et al., 2014). 20 species. Rumohra Raddi, Opusc. Sci. 3: 290. 1819. Type: Rumohra aspidioides Raddi ( Polypodium adiantiforme G.Forst.). Circumscription sensu Rakotondrainibe (2010), Sundue et al. (2013), and Labiak et al. (2014). Monophyletic (Labiak et al., 2014). Eight species. Teratophyllum Mett. ex Kuhn, Ann Mus. Bot. Lugduno-Batavi 4(10): 296. 1869. Lectotype (designated by Holttum, Gard. Bull. Straits Settlem. 5: 277. 1932): Teratophyllum aculeatum (Blume) Mett. ex Kuhn ( Lomaria aculeata Blume). Circumscription sensu Moran et al. (2010a). Monophyletic (Moran et al., 2010a). 13 species. Subfamily Dryopteridoideae Link, Fil. Spec.: 116. 1841 (Dryopterideae). Circumscription sensu Liu et al. (2016). Monophyletic (Liu et al., 2016). Six genera and an estimated 1128 species. J. Syst. Evol. 54 (6): 563–603, 2016 588 The Pteridophyte Phylogeny Group Arachniodes Blume, Enum. Pl. Javae 2: 241. 1828. Type: Arachniodes aspidioides Blume. Circumscription sensu He et al. (2013) and Liu et al. (2016). Includes: Byrsopteris C.V.Morton; Leptorumohra (H.Ito) H.Ito; Lithostegia Ching; Phanerophlebiopsis Ching. Assumed to be monophyletic (Liu et al., 2016). 60 species. Ctenitis (C.Chr.) C.Chr., Man. Pteridol.: 544. 1938. Lectotype (designated by Ching, Bull. Fan Mem. Inst. Biol. Bot. 8(5): 275. 1938): Ctenitis distans (Brack.) Ching ( Lastrea distans Brack.). Circumscription sensu Wang et al. (2014), Liu et al. (2016), and Duan et al. (2017). Includes: Ataxipteris Holttum; Pseudotectaria Tardieu. Monophyletic (Wang et al., 2014; Hennequin et al., pers. comm.). About 125 species. Cyrtomium C.Presl, Tent. Pterid.: 86, pl.2, f.26. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 204. 1875): Cyrtomium falcatum (L.f.) C.Presl ( Polypodium falcatum L.f.). Circumscription sensu Lu et al. (2007), Zhang et al. (2013b), and Liu et al. (2016). Assumed to be monophyletic (Lu et al., 2007). 35 species. Dryopteris Adans., Fam. Pl. 2: 20. 1763. Type: Dryopteris filix-mas (L.) Schott ( Polypodium filix-mas L.). Circumscription sensu Zhang & Zhang (2012), Zhang et al. (2012), and Zhang et al. (2013b). Includes: Acrophorus C.Presl; Acrorumohra (H.Ito) H.Ito; Adenoderris J.Sm, in part; Arthrobotrys (C.Presl) Lindl.; Diacalpe Blume; Dichasium (A.Braun) F
ee; Diclisodon T. Moore; Dryopsis Holttum & P.J.Edwards; Filix S
eg.; Lophodium Newman; Nephrodium Marthe ex Michx.; Nothoperanema (Tagawa) Ching; Peranema D.Don; Pycnopteris T.Moore; Revwattsia D.L.Jones; Stenolepia Alderw. Monophyletic (Zhang et al., 2012; Sessa et al., 2012; Zhang & Zhang, 2012; McKeown et al., 2012; Kuo et al., 2016a). About 400 species. Phanerophlebia C.Presl, Tent. Pterid.: 84, pl.2, f.19. 1836. Type: Phanerophlebia nobilis (Schltdl. & Cham.) C.Presl ( Aspidium nobile Schltdl. & Cham.). Circumscription sensu Yatskievych et al. (1988) and Liu et al. (2016). Includes: Amblia C. Presl. Monophyletic (Yatskievych et al., 1988). Eight species. Polystichum Roth, Tent. Fl. Germ. 3(1): 31, 69–70. 1800. Lectotype (designated by J.Sm., Hist. Fil.: 217. 1875): Polystichum aculeatum (L.) Roth ( Polypodium aculeatum L.). Circumscription
chon et al. sensu Zhang et al. (2013b) and Le Pe (2016). Includes: Acropelta T.Nakai; Adenoderris J.Sm, in part; Cyrtogonellum Ching; Cyrtomidic
v.; Hypopeltis tyum Ching; Hemesteum H.Le
e; Michx.; Papuapteris C.Chr.; Plecosorus Fe J. Syst. Evol. 54 (6): 563–603, 2016 Ptilopteris Hance; Sorolepidium Christ. Monophyletic (Le P
echon et al., 2016). About 500 species. Subfamily placement uncertain. Aenigmopteris Holttum, Blumea 30: 3. 1984. Type: Aenigmopteris dubia (Copel.) Holttum ( Dryopteris dubia Copel.). Circumscription sensu Kramer et al. (1990). No molecular data available. Five species. Dryopolystichum Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 125. 1947. Type: Dryopolystichum phaeostigma (Ces.) Copel. ( Aspidium phaeostigma Ces.). Circumscription sensu Christenhusz et al. (2011). Monotypic (Christenhusz et al., 2011). 46. Family Nephrolepidaceae Pic.Serm., Webbia 29(1): 8–11. 1975. Circumscription sensu Hovenkamp & Miyamoto (2012). Family consists of a single genus and an estimated 19 species. Nephrolepis Schott, Gen. Fil.: 1, pl.3. 1834. Lectotype (designated by J.Sm., Hist. Fil. 226. 1875): Nephrolepis exaltata (L.) Schott ( Polypodium exaltatum L.). Circumscription sensu Hennequin et al. (2010). Monophyletic (Hennequin et al., 2010; Zhang et al., 2016). 19 species. 47. Family Lomariopsidaceae Alston, Taxon 5(2): 25. 1956. Circumscription sensu Christenhusz et al. (2013). Monophyletic (Christenhusz et al., 2013; Zhang et al., 2016). Four genera and an estimated 69 species. Cyclopeltis J.Sm., Bot. Mag. 72: 36. 1846. Type: Cyclopeltis semicordata (Sw.) J.Sm. ( Polypodium semicordatum Sw.). Circumscription sensu Kramer et al. (1990). Monophyletic (Zhang & Zhang, 2015). Six species. Dracoglossum Christenh., Thaiszia 17(1–2): 3–5. 2007. Type: Dracoglossum plantagineum (Jacq.) Christenh. ( Polypodium plantagineum Jacq.). Circumscription sensu Christenhusz (2007). Monophyletic (Zhang et al., 2016). Two species.
e, Me
m. Foug., 2. Hist. Acrostich.: Lomariopsis Fe 10. 1845. Lectotype (designated by J.Sm., Hist.
e ( Fil.: 140. 1875): Lomariopsis sorbifolia (L.) Fe Acrostichum sorbifolium L.). Circumscription sensu Rouhan et al. (2007) and Moran (2000). Monophyletic (Rouhan et al., 2007; Li et al., 2009). About 60 species. Thysanosoria A.Gepp, Fl. Arfak Mts. (Gibbs): 193– 194, pl.4. 1917. Type: Thysanosoria dimorphophylla A.Gepp. Circumscription sensu Kramer (1990h). Genus has yet to be included in a phylogenetic analysis. Monotypic. www.jse.ac.cn PPG I 48. Family Tectariaceae Panigrahi, J. Orissa Bot. Soc. 8: 41. 1986. Circumscription sensu Zhang et al. (2016). Monophyletic (Zhang et al., 2016). Seven genera and an estimated 250 species. Arthropteris J.Sm. ex Hook.f., Fl. Nov.-Zel. 2: 43. 1855. Type: Arthropteris tenella (G.Forst.) J.Sm. ex Hook.f. ( Polypodium tenellum G.Forst.). Circumscription sensu Liu et al. (2013). Includes: Psammiosorus C.Chr. Monophyletic (Liu et al., 2013; Zhang et al., 2016). About 15 species. Draconopteris Li Bing Zhang & Liang Zhang, Taxon 65(4): 732, f.1a–b, f.3a–d. 2016. Type: Draconopteris draconoptera (D.C.Eaton) Li Bing Zhang & Liang Zhang ( Aspidium draconopterum D.C.Eaton). Circumscription sensu Zhang et al. (2016). Monotypic. Hypoderris R.Br. in Wall., Pl. Asiat. Rar. (Wallich) 1: 16. 1829. Lectotype (designated by J.Sm. in Hook. Gen. Fil.: t.1. 1838.): Hypoderris brownii J.Sm. Circumscription sensu Moran et al. (2014). Monophyletic (Moran et al., 2014; Zhang et al., 2016). Three species. Malaifilix Li Bing Zhang & Schuettp., Taxon 65(4): 733, f.1e–f, f.3e–f. 2016. Type: Malaifilix grandidentata (Ces.) Li Bing Zhang & Schuettp. ( Polypodium dilatatum var. grandidentatum Ces.). Circumscription sensu Zhang et al. (2016). Presumably monotypic. Pteridrys C.Chr. & Ching, Bull. Fan Mem. Inst. Biol. Bot. 5(3): 129. 1934. Type: Pteridrys syrmatica (Willd.) C.Chr. & Ching ( Aspidium syrmaticum Willd.). Circumscription sensu Christensen & Ching (1934). Monophyletic (Zhang et al., 2016). About 10 species. Tectaria Cav., Anales Hist. Nat. 1(2): 115. 1799. Type: Tectaria trifoliata (L.) Cav. ( Polypodium trifoliatum L.). Circumscription sensu Zhang et al. (2016). Includes: Amphiblestra C.Presl; Aspidium
e; Sw.; Bathmium C.Presl ex Link; Camptodium Fe Chlamydogramme Holttum; Cionidium T.Moore; Ctenitopsis Ching ex Tardieu & C.Chr.; Dictyox
e ex J.Sm.; Fadyenia iphium Hook.; Dryomenis Fe Hook.; Grammatosorus Regel; Hemigramma Christ; Heterogonium C.Presl; Lenda Koidz.; Luerssenia Kuhn ex Luerssen; Microbrochis C.Presl;
e; Podopeltis Fe
e; Polydictyum Phlebiogonium Fe C.Presl; Psomiocarpa C.Presl; Quercifilix Copel.; Sagenia C.Presl; Stenosemia C.Presl; Tectaridium Copel. Monophyletic (Zhang et al., 2016), but molecular data is lacking for Amphiblestra, Camptodium, Chlamydogramme, Dryomenis, Grammatosorus, Lenda, Luerssenia, Microbrochis, Phlebiogonium, and Polydictyum (tentatively included here based on morphology). About 200 species. www.jse.ac.cn 589 Triplophyllum Holttum, Kew Bull. 41(2): 239. 1986. Type: Triplophyllym protensum (Afzel. ex Sw.) Holttum ( Aspidium protensum Afzel. ex Sw.). Circumscription sensu Holttum (1986). Monophyletic (Prado & Moran, 2008). About 20 species. 49. Family Oleandraceae Ching ex Pic.Serm., Webbia 20 (2): 745. 1965. Circumscription sensu Smith et al. (2006b). Family consists of a single genus and an estimated 15 species. Oleandra Cav., Anales Hist. Nat. 1(2): 115. 1799. Type: Oleandra neriiformis Cav. Circumscription sensu Kramer (1990n). Monophyletic (Zhang & Zhang, 2015). At least 15 species. 50. Family Davalliaceae M.R.Schomb., Reis. Br.-Guiana 3: 883. 1848. Circumscription sensu Smith et al. (2006b). Family consists of a single genus and an estimated 65 species.
m. Acad. Roy. Sci. (Turin) 5: 414, Davallia Sm., Me
e, pl. 9, f. 6. 1793. Lectotype (designated by Fe
m. Foug., 5. Gen. Filic.: 328. 1852): Davallia Me canariensis (L.) Sm. ( Trichomanes canariense L.). Circumscription sensu Tsutsumi et al. (2016). Includes: Araiostegia Copel.; Araiostegiella M.Kato & Tsutsumi; Davallodes (Copel.) Copel.; Humata
e; Wibelia Bernh. MonophyCav.; Scyphularia Fe letic (Tsutsumi & Kato, 2006; Tsutsumi et al., 2016). About 65 species. 51. Family Polypodiaceae J.Presl & C.Presl, Delic. Prag.: 159. 1822. Circumscription sensu Smith et al. (2006b). Includes: Loxogrammaceae Ching ex Pic.Serm.; Grammitidaceae Newman; Platyceriaceae Ching; Drynariaceae Ching. Monophyletic (Schuettpelz & Pryer, 2007). Six subfamilies, 65 genera, and an estimated 1652 species. Subfamily Loxogrammoideae H.Schneid. in Christenhusz et al., Phytotaxa 19: 18. 2011. Circumscription equivalent to “loxogrammoid clade“ in Kreier & Schneider (2006a). Monophyletic (Kreier & Schneider, 2006a). Two genera and an estimated 32 species. Dictymia J.Sm., Companion Bot. Mag. 72: 16. 1846. Type: Dictymia attenuata J.Sm. Circumscription sensu Kreier & Schneider (2006a). Monophyletic (Kreier & Schneider, 2006a). No published generic revision available. About two species. Loxogramme (Blume) C.Presl, Tent. Pterid.: 214–215, pl.9, f.8. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 156. 1875): Loxogramme lanceolata (Sw.) C.Presl ( Grammitis lanceolata Sw.). Circumscription sensu Kreier & Schneider (2006a). Includes: Anarthropteris Copel. Monophyletic (Kreier & Schneider, 2006a). No published generic revision available. 30 species. J. Syst. Evol. 54 (6): 563–603, 2016 590 The Pteridophyte Phylogeny Group Subfamily Platycerioideae B.K.Nayar, Taxon 19: 233. 1970. Circumscription sensu Hennipman et al. (1990). Monophyletic (Kreier & Schneider, 2006b). Two genera and an estimated 70 species. Platycerium Desv., M
em. Soc. Linn. Paris 6(3): 213. 1827. Lectotype (designated by Pfeiff., Nomencl. Bot. 2: 746. 1873): Platycerium alcicorne Desv. Circumscription sensu Hennipman et al. (1990). Monophyletic (Kreier & Schneider, 2006b). 18 species. Pyrrosia Mirb., Hist. Nat. Veg. 3: 471. 1803. Type: Pyrrosia chinensis Desv. Circumscription sensu Hennipman et al. (1990). Includes: Drymoglossum C.Presl. Assumed to be monophyletic (Kreier & Schneider, 2006b), but paraphyly relative to Platycerium has not yet been definitively excluded. At least 52 species. Subfamily Drynarioideae Crabbe, Jermy & Mickel, Fern Gaz. 11: 156. 1975. Circumscription sensu Christenhusz et al. (2011). Monophyletic (Schneider et al., 2008). Six genera and an estimated 148 species. Aglaomorpha Schott, Gen. Fil.: t.19. 1836. Type: Aglaomorpha meyeniana Schott. Circumscription newly proposed in this classification. Includes: Christiopteris Copel., Drynaria J.Sm., Dryostachyum J.Sm., Merinthosorus Copel., Photinopteris J.Sm., Pseudodrynaria C.Chr., Thayeria Copel. Monophyletic (Schneider et al., 2008). 50 species. Arthromeris (T.Moore) J.Sm., Hist. Fil.: 110. 1875. Type: Arthromeris wallichiana (Spreng.) Ching ( Polypodium wallichianum Spreng.). Circumscription sensu Hennipman et al. (1990). Assumed to be monophyletic (Schneider et al., 2008), but comprehensive analysis is lacking to date. Perhaps a few more species, if a narrower species concept is used. Eight species. Gymnogrammitis Griff., Icon. Pl. Asiat. 2: t.129, f.1. 1849. Type: Gymnogrammitis dareiformis (Hook.) Ching ex Tardieu & C.Chr. ( Polypodium dareiforme Hook.). Circumscription sensu Kramer (1990c). Monotypic. Paraselliguea Hovenkamp, Blumea 45(2): 376. 2000. Type: Paraselliguea leucophora (Baker) Hovenkamp ( Polypodium leucophorum Baker). Circumscription sensu Hovenkamp (1997). Monotypic. Polypodiopteris C.F.Reed, Amer. Fern J. 38: 87. 1948. Type: Polypodiopteris proavita (Copel.) C.F. Reed ( Polypodium proavitum Copel.). Circumscription sensu Hennipman et al. (1990). Assumed to be monophyletic, but only one species has been sampled to date. Three species. J. Syst. Evol. 54 (6): 563–603, 2016 Selliguea Bory, Dict. Class. Hist. Nat. 6: 587–588. 1824. Type: Selliguea feei Bory. Circumscription sensu Schneider et al. (2008). Includes: Crypsinus C.Presl; Himalayopteris W.Shao & S.G.Lu; Phymatopsis J.Sm.; Pichisermollodes Fraser-Jenk. & Challis. Not monophyletic as circumscribed (Schneider et al., 2008; Li et al., 2012a); Selliguea appears to be polyphyletic relative to Arthromeris, Gymnogrammitis, and Polypodiopteris, but additional study is needed. At least 85 species. Subfamily Microsoroideae B.K.Nayar, Taxon 19(2): 233. 1970. Circumscription equivalent to “microsoroids” in Kreier et al. (2008b). Monophyletic (Schneider et al., 2004c). 12 genera and an estimated 183 species. Goniophlebium (Blume) C.Presl, Tent. Pterid.: 185, € dlpl.7, f.13–14. 1836. Lectotype (designated by Ro Linder, Zijlstra & Tryon, Taxon 39(1): 105. 1990): Goniophlebium subauriculatum (Blume) C.Presl ( Polypodium subauriculatum Blume). Circumscription sensu Kreier et al. (2008b). Includes: Metapolypodium Ching; Polypodiastrum Ching; Polypodiodes Ching. Monophyletic (Kreier et al., 2008b). 25 species. Lecanopteris Reinw. ex Blume, Enum. Pl. Javae 2: 120. 1828. Type: Lecanopteris carnosa (Reinw.) Blume ( Onychium carnosum Reinw.). Circumscription sensu Hennipman et al. (1990). Monophyletic (Kreier et al., 2008b). 13 species. Lemmaphyllum C.Presl, Abh. Koenigl. B€ ohm. Ges. Wiss. Ser. 5 6: 517–518. 1851. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 189. 1947): Lemmaphyllum spathulatum C.Presl. Circumscription sensu Wang et al. (2010). Includes: Caobangia A.R.Sm. & X.C.Zhang; Lepidogrammitis Ching; Weatherbya Copel. Monophyletic (Wang et al., 2010). At least two species. Lepidomicrosorium Ching & K.H.Shing, Bot. Res. Academia Sinica 1(1): 1–14, pl.1–5. 1983. Type: Lepidomicrosorium subhastatum (Baker) Ching ( Polypodium subhastatum Baker). Circumscription sensu Wang et al. (2010). Apparently monotypic (Wang et al., 2010). Lepisorus (J.Sm.) Ching, Bull. Fan Mem. Inst. Biol. Bot. 4(3): 47, 56–58. 1933. Type: Lepisorus nudus (Hook.) Ching ( Pleopeltis nuda Hook.). Circumscription sensu Wang et al. (2010). Includes: Belvisia Mirb.; Drymotaenium Makino. Monophyletic (Wang et al., 2010). 80 species. Leptochilus Kaulf., Enum. Filic.: 147, pl.1, f.10. 1824. Type: Leptochilus axillaris (Cav.) Kaulf. ( Acrostichum axillare Cav.). Circumscription sensu Kreier et al. (2008b). Includes: Colysis C.Presl; Kontumia www.jse.ac.cn PPG I S.K.Wu & K.L.Phan. Monophyletic (Kreier et al., 2008b). 10 species. Microsorum Link, Hort. Berol. 2: 110. 1833. Type: Microsorum irregulare Link. Circumscription sensu Kreier et al. (2008b). Includes: Dendroconche Copel.; Kaulinia B.K.Nayar; Phymatosorus Pic. Serm. Not monophyletic as circumscribed (Kreier et al., 2008b); Microsorum is apparently paraphyletic relative to Lecanopteris, Leptochilus, Neocheiropteris, Neolepisorus, and Tricholepidium, but further study is needed. 40 species. Neocheiropteris Christ, Bull. Soc. Bot. France 52
m. 1): 21. 1905. Type: Neocheiropteris palma(Me topedata (Baker) Christ ( Polypodium palmatopedatum Baker). Circumscription sensu Wang et al. (2010). Monophyletic (Kreier et al., 2008b; Wang et al., 2010; Du & Cheng, 2011). Two species. Neolepisorus Ching, Bull. Fan Mem. Inst. Biol. Bot. 10(1): 11–12. 1940. Type: Neolepisorus ensatus (Thunb.) Ching ( Polypodium ensatum Thunb.). Circumscription sensu Wang et al. (2010). Monophyletic (Wang et al., 2010). The generic affinities have not been evaluated for all species possibly belonging here. At least five species. Paragramma (Blume) T.Moore, Index Filic. xxxii. 1857. Lectotype (designated by J.Sm., Hist. Fil.: 114. 1875): Paragramma longifolia (Blume) T. Moore ( Grammitis longifolia Blume). Circumscription sensu Wang et al. (2010). Only the type species has been sampled to date; the second species is highly distinct and may ultimately be resolved elsewhere. Perhaps two species. 591 Le
on (1995), Schneider et al. (2004c), and Kreier et al. (2007). Includes: Hyalotricha Copel.; Hyalotrichopteris W.H.Wagner. Monophyletic (Schneider et al., 2004c). About 50 species. Microgramma C.Presl, Tent. Pterid.: 213. 1836. Type: Microgramma persicariifolia (Schrad.) C. Presl ( Polypodium persicariifolium Schrad.). Circumscription sensu Salino et al. (2008). Includes: Solanopteris Copel. Monophyletic (Salino et al., 2008). A generic revision is lacking. About 30 species. Niphidium J.Sm., Hist. Fil.: 99. 1875. Type: Niphidium americanum (Hook.) J.Sm. ( Polypodium americanum Hook.). Circumscription sensu Hennipman et al. (1990). Monophyletic (Salino et al., 2008). 10 species. Pecluma M.G.Price, Amer. Fern J. 73(4): 109. 1983. Type: Pecluma pectinata (L.) M.G.Price ( Polypodium pectinatum L.). Circumscription sensu Assis et al. (2016). Monophyletic (Assis et al., 2016). 40 species. Phlebodium (R.Br.) J.Sm., J. Bot. (Hooker) 4: 58. 1841. Lectotype (designated by J.Sm., Hist. Fil.: 94. 1875): Phlebodium aureum (L.) J.Sm. ( Polypodium aureum L.). Circumscription sensu Mickel & Smith (2004). Monophyletic (Assis et al., 2016). Four species. Thylacopteris Kunze ex J.Sm., Hist. Fil.: 87. 1875. Type: Thylacopteris papillosa (Blume) J.Sm. ( Polypodium papillosum Blume). Circumscription sensu Hennipman et al. (1990). Assumed to be monophyletic, but only the type species has been sampled to date. Two species. Pleopeltis Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5: 211. 1810. Type: Pleopeltis angusta Humb. & Bonpl. ex Willd. Circumscription sensu Smith & Tejero-D
ıez (2014). Includes: Cheilogramma Maxon; Dicranoglossum J.Sm.; Eschatogramme Trevis.; Lepicystis (J.Sm.) J.Sm.; Marginaria Bory; Marginariopsis C.Chr.; Microphlebodium L.D. G
omez; Neurodium F
ee; Paltonium C.Presl; Pseudocolysis L.D.G
omez. Monophyletic (Otto et al., 2009). In need of a generic revision. About 90 species. Tricholepidium Ching, Acta Phytotax. Geobot. 29 (1–5): 41. 1978. Type: Tricholepidium normale (D. Don) Ching ( Polypodium normale D.Don). Circumscription sensu Wang et al. (2010). Monotypic (Wang et al., 2010), but in need of further evaluation. Pleurosoriopsis Fomin, Izv. Kievsk. Bot. Sada 11: 8. 1930. Type: Pleurosoriopsis makinoi (Maxim. ex Makino) Fomin ( Gymnogramma makinoi Maxim. ex Makino). Circumscription sensu Hennipman et al. (1990). Monotypic (Assis et al., 2016). Subfamily Polypodioideae Sweet, Hort. Brit.: 460. 1826. Newly circumscribed in this classification. Paraphyletic relative to the Grammitidoideae (Schneider et al., 2004c). Nine genera and an estimated 305 species. Polypodium L., Sp. Pl. 2: 1082. 1753. Lectotype (designated by J.Sm., Hist. Fil.: 88. 1875): Polypodium vulgare L. Circumscription sensu Sigel et al. (2014) and Assis et al. (2016). Presumably 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. 1875): Campyloneurum repens (Aubl.) C.Presl ( Polypodium repens Aubl.). Circumscription sensu www.jse.ac.cn Serpocaulon A.R.Sm., Taxon 55(4): 924–927, f. 34. 2006. Type: Serpocaulon loriceum (L.) A.R.Sm. ( Polypodium loriceum L.). Circumscription J. Syst. Evol. 54 (6): 563–603, 2016 592 The Pteridophyte Phylogeny Group sensu Smith et al. (2006a). Monophyletic (Smith et al., 2006a; Kreier et al., 2008a). About 40 species. Subfamily Grammitidoideae Parris & Sundue, this classification (see below). Circumscription equivalent to “grammitids” in Sundue et al. (2014). Monophyletic (Ranker et al., 2004; Schneider et al., 2004c; Sundue et al., 2014). 33 genera and an estimated 911 species. Acrosorus Copel., Philipp. J. Sci. 1(Suppl. 2): 158– 159. 1906. Lectotype (designated by Copel., Univ. Calif. Publ. Bot. 16: 108. 1929): Acrosorus exaltatus (Copel.) Copel. ( Davallia exaltata Copel.). Circumscription sensu Parris (1990) and Sundue et al. (2014). Assumed to be monophyletic, but only one species sampled to date (Sundue et al., 2014). Nine species. Adenophorus Gaudich., Ann. Sci. Nat. (Paris) 3: 508. 1824. Lectotype (designated by J.Sm., Hist. Fil.: 187. 1875): Adenophorus tripinnatifidus Gaudich. Circumscription sensu Ranker (2008) and Sundue et al. (2014). Includes: Amphoradenium Desv. Monophyletic (Sundue et al., 2014). 10 species. Alansmia M.Kessler, Moguel, Sundue & Labiak, Brittonia 63(2): 238. 2011. Type: Alansmia lanigera (Desv.) Moguel & M.Kessler ( Polypodium lanigerum Desv.). Circumscription sensu Kessler et al. (2011) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 26 species. Archigrammitis Parris, Fern Gaz. 19(4): 135–136. 2013. Type: Archigrammitis friderici-et-pauli (Christ) Parris ( Polypodium friderici-et-pauli Christ). Circumscription sensu Parris (2013). No molecular data available. Seven species. Ascogrammitis Sundue, Brittonia 62(4): 361. 2010. Type: Ascogrammitis athyrioides (Hook.) Sundue ( Polypodium athyrioides Hook.). Circumscription sensu Sundue (2010) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 17 species. Calymmodon C.Presl, Tent. Pterid.: 203–204, pl.9, f.1. 1836. Type: Calymmodon cucullatus (Nees & Blume) C.Presl ( Polypodium cucullatum Nees & Blume). Circumscription sensu Parris (1990) and
e. Sundue et al. (2014). Includes: Plectopteris Fe Monophyletic (Sundue et al., 2014). 65 species. Ceradenia L.E.Bishop, Amer. Fern J. 78(1): 2. 1988. Type: Ceradenia curvata (Sw.) L.E.Bishop ( Polypodium curvatum Sw.). Circumscription sensu Bishop (1988) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). About 73 species. J. Syst. Evol. 54 (6): 563–603, 2016 Chrysogrammitis Parris, Kew Bull. 3(4): 909. 1998. Type: Chrysogrammitis glandulosa (J.Sm.) Parris ( Ctenopteris glandulosa J.Sm.). Circumscription sensu Parris (1998) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). Two species. Cochlidium Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 36. 1820. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 213. 1947): Cochlidium graminoides (Sw.) Kaulf. ( Acrostichum graminoides Sw.). Circumscription sensu Bishop (1978) and Sundue et al. (2014). Includes: Xiphopteris Kaulf.; Micropteris Desv.; Pleurogramme Blume. Monophyletic, but nested within Grammitis (Sundue et al., 2014). Grammitis s.s. was found to be paraphyletic with regards to Cochlidium in Sundue et al. (2014), but the type, Grammitis marginella, has not been sampled. We prefer to maintain both genera as they are circumscribed until additional data becomes available. 18 species. Ctenopterella Parris, Gard. Bull. Singapore 58(2): 234. 2007. Type: Ctenopterella blechnoides (Grev.) Parris ( Grammitis blechnoides Grev.). Circumscription sensu Parris (2007) and Sundue et al. (2014). Only one species sampled to date (Sundue et al., 2014). 24 species. Dasygrammitis Parris, Gard. Bull. Singapore 58(2): 238. 2007. Type: Dasygrammitis mollicoma (Nees & Blume) Parris ( Polypodium mollicomum Nees & Blume). Circumscription sensu Parris (2007) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 12 species. Enterosora Baker, Timehri 5: 218. 1886. Type: Enterosora campbellii Baker. Circumscription sensu Bishop & Smith (1992) pro parte and Sundue et al. (2014). Monophyletic, but nested within Zygophlebia (Sundue et al., 2014). This could be resolved by placing Zygophlebia in synonymy. However, this would be premature since the type of Enterosora has not yet been sampled and its placement is not certain. 11 species. Galactodenia Sundue & Labiak, Syst. Bot. 37(2): 340. 2012. Type: Galactodenia delicatula (M. Martens & Galeotti) Sundue & Labiak ( Polypodium delicatulum M.Martens & Galeotti). Circumscription sensu Sundue et al. (2012, 2014). Monophyletic (Sundue et al., 2014). Five species. Grammitis Sw., J. Bot. (Schrader) 2: 3, 17. 1801. Lectotype (designated by C.Chr., Index Filic. xlix. 1906): Grammitis marginella (Sw.) Sw. ( Polypodium marginellum Sw.). Circumscription sensu Bishop (1977) pro parte and Sundue et al. (2014). www.jse.ac.cn PPG I Grammitis s.s. is paraphyletic relative to Cochlidium (Sundue et al., 2014), but the type, G. marginella, has not been sampled. We prefer to maintain both genera as they are circumscribed until additional data become available. 43 species. Lellingeria A.R.Sm. & R.C.Moran, Amer. Fern J. 81 (3): 76. 1991. Type: Lellingeria apiculata (Kunze ex Klotzsch) A.R.Sm. & R.C.Moran ( Polypodium apiculatum Kunze ex Klotzsch). Circumscription sensu Labiak (2013) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 49 species. Leucotrichum Labiak, Taxon 59(3): 915. 2010. Type: Leucotrichum organense (Gardner) Labiak ( Grammitis organensis Gardner). Circumscription sensu Labiak et al. (2010), Rouhan et al. (2012), and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). Six species. Lomaphlebia J.Sm., Hist. Fil.: 182. 1875. Type: Lomaphlebia linearis (Sw.) J.Sm. ( Grammitis linearis Sw.). Circumscription sensu Sundue et al. (2014). Assumed to be monophyletic, but only one species sampled to date (Sundue et al., 2014). Two species. Luisma M.T.Murillo & A.R.Sm., Novon 13(3): 313– 316, f. 1. 2003. Type: Luisma bivascularis M.T. Murillo & A.R.Sm. Circumscription sensu Murillo & Smith (2003). Monotypic. Melpomene A.R.Sm. & R.C.Moran, Novon 2(4): 426. 1992. Type: Melpomene moniliformis (Lag. ex Sw.) A.R.Sm. & R.C.Moran ( Polypodium moniliforme Lag. ex Sw.). Circumscription sensu Lehnert (2013) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 29 species. Micropolypodium Hayata, Bot. Mag. (Tokyo) 42 (449): 341. 1928. Type: Micropolypodium pseudotrichomanoides (Hayata) Hayata ( Polypodium pseudotrichomanoides Hayata). Circumscription sensu Smith (1992) pro parte, Hirai et al. (2011), and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). Three species. Moranopteris R.Y.Hirai & J.Prado, Taxon 60(4): 1127. 2011. Type: Moranopteris basiattenuata (Jenman) R.Y.Hirai & J.Prado ( Polypodium basiattenuatum Jenman). Circumscription sensu Hirai et al. (2011) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 31 species. Mycopteris Sundue, Brittonia 66(2): 175, f.1–2. 2013. Type: Mycopteris taxifolia (L.) Sundue ( Polypodium taxifolium L.). Circumscription sensu Sundue (2013) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 21 species. www.jse.ac.cn 593 Notogrammitis Parris, New Zealand J. Bot. 50(4): 465. 2012. Type: Notogrammitis billardierei (Willd.) Parris ( Grammitis billardierei Willd.). Circumscription sensu Perrie and Parris (2012). Monophyletic (Sundue et al., 2014). 16 species. Oreogrammitis Copel., Philipp. J. Sci. 12: 64. 1917. Type: Oreogrammitis clemensiae Copel. Circumscription sensu Parris (2007) and Sundue et al. (2014). Not monophyletic and in need of redefinition (Sundue et al., 2014). Themelium is monophyletic in Sundue et al. (2014), but nested within a clade containing polyphyletic Oreogrammitis and Radiogrammitis. The types of Oreogrammitis and Themelium remain to be sampled, and thus we recommend maintaining these genera pending further research. 156 species. Prosaptia C.Presl, Tent. Pterid.: 165–166, pl.6, f.19, 25. 1836. Lectotype (designated by J.Sm., Hist. Fil.: 86. 1875): Prosaptia contigua (G.Forst.) C. Presl ( Trichomanes contiguum G.Forst.). Circumscription sensu Parris (2010a, 2010b) and Sundue et al. (2014). Includes: Ctenopteris Blume
e. Monophyletic (Sunex Kunze; Cryptosorus Fe due et al., 2014). 87 species. Radiogrammitis Parris, Gard. Bull. Singapore 58 (2): 240. 2007. Type: Radiogrammitis setigera (Blume) Parris ( Polypodium setigerum Blume). Circumscription sensu Parris (2007) and Sundue et al. (2014). Not monophyletic and in need of redefinition (Sundue et al., 2014). Themelium is monophyletic in Sundue et al. (2014), but nested within a clade containing polyphyletic Oreogrammitis and Radiogrammitis. The types of Oreogrammitis and Themelium remain to be sampled, and thus we recommend maintaining these genera pending further research. 36 species. Scleroglossum Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 2(7): 37–39. 1912. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 213. 1947): Scleroglossum pusillum (Blume) Alderw. ( Vittaria pusilla Blume). Circumscription sensu Parris (1990) and Sundue et al. (2014). Includes: Nematopteris Alderw. Monophyletic (Sundue et al., 2014). Seven species. Stenogrammitis Labiak, Brittonia 63(1): 141, f.1A– M, 2A–F. 2011. Type: Stenogrammitis myosuroides (Sw.) Labiak ( Polypodium myosuroides Sw.). Circumscription sensu Labiak (2011) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 31 species. Terpsichore A.R.Sm., Novon 3(4): 479. 1993. Type: Terpsichore asplenifolia (L.) A.R.Sm. ( Polypodium asplenifolium L.). Circumscription sensu J. Syst. Evol. 54 (6): 563–603, 2016 594 The Pteridophyte Phylogeny Group Smith (1993) pro parte and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 12 species. Themelium (T.Moore) Parris, Kew Bull. 52(3): 737. 1997. Type: Themelium tenuisectum (Blume) Parris ( Polypodium tenuisectum Blume). Circumscription sensu Parris (1997, 2004, 2010b) and Sundue et al. (2014). Themelium is monophyletic in (Sundue et al., 2014), but nested within a clade containing polyphyletic Oreogrammitis and Radiogrammitis. The types of Oreogrammitis and Themelium remain to be sampled, and thus we recommend maintaining these genera pending further research. 29 species. Tomophyllum (E.Fourn.) Parris, Gard. Bull. Singapore 58(2): 245. 2007. Lectotype (designated by Parris, Gard. Bull. Singapore 58(2): 245. 2007): Tomophyllum subsecundodissectum (Zoll.) Parris ( Polypodium subsecundodissectum Zoll.). Circumscription sensu Parris (2007) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 42 species. Xiphopterella Parris, Gard. Bull. Singapore 58(2): 249. 2007. Type: Xiphopterella hieronymusii (C. Chr.) Parris ( Polypodium hieronymusii C.Chr.). Circumscription sensu Parris (2007) and Sundue et al. (2014). Monophyletic (Sundue et al., 2014). 11 species. Zygophlebia L.E.Bishop, Amer. Fern J. 79(3): 107. 1989. Type: Zygophlebia sectifrons (Kunze ex Mett.) L.E.Bishop ( Polypodium sectifrons Kunze ex Mett.). Circumscription sensu Bishop (1989) and Sundue et al. (2014). Paraphyletic relative to Enterosora (Sundue et al., 2014). This could be resolved by placing Zygophlebia in synonymy. However, this would be premature since the type of Enterosora has not yet been sampled and its placement is not certain. 20 species. Subfamily placement uncertain. Synammia C.Presl, Tent. Pterid.: 212, t.9, f.11. 1836. Lectotype (designated by Copel., Ann. Cryptog. Phytopathol. 5 [Gen. Fil.]: 212. 1947): Synammia triloba (Cav.) C.Presl. Circumscription sensu Schneider et al. (2006). Monophyletic (Schneider et al., 2006). Three species. Nomenclatural novelties Aspleniineae H.Schneid. & C.J.Rothf., stat. nov.  Aspleniaceae Newman, Hist. Brit. Ferns 6. 1840. Type: Asplenium L. (1753)—Polypodiales. Claytosmunda (Y.Yatabe, N.Murak. & K.Iwats.) Metzgar & Rouhan, stat. nov.  Osmunda subg. Claytosmunda Y.Yatabe, J. Syst. Evol. 54 (6): 563–603, 2016 N.Murak. & K.Iwats., Acta Phytotax. Geobot. 56(2): 127–128. 2005. Type: Claytosmunda claytoniana (L.) Metzgar & Rouhan ( Osmunda claytoniana L.). Claytosmunda claytoniana (L.) Metzgar & Rouhan, comb. nov.  Osmunda claytoniana L., Sp. Pl. 2: 1066. 1753. Dennstaedtiineae Schwartsb. & Hovenkamp, subord. nov. Type: Dennstaedtia Bernh. (1801) [1800]. Type: Dennstaedtiaceae—Polypodiales. Rhizome generally dorsiventral and long-creeping, covered with hairs or proto-scales (true scales absent); sori marginal or sub-marginal, generally protected by marginal pseudo-indusia or by laminar true indusia (sometimes both are present), rarely fully exindusiate. Grammitidoideae Parris & Sundue, stat. nov.  Grammitidaceae Newman, Hist. Brit. Ferns: 7. 1840, as “Grammitideae”. Type: Grammitis Sw. (1801) [1800]—Polypodiaceae. Lindsaeineae Lehtonen & Tuomisto, subord. nov. Type: Lindsaea Dryand. ex J.Sm. (1793), as Lindsaeaceae— Polypodiales. Rhizome short- to long-creeping (rarely ascending), covered with non-clathrate scales or rarely hairs; petioles with one or rarely two (or several fusing into two in the upper part of stipe) vascular bundles; sori marginal to sub-marginal, generally protected by laminar true indusia or rarely by marginal pseudo-indusia or both. Mankyuoideae J.R.Grant & B.Dauphin, subfam. nov. Type: Mankyua B.Y.Sun, M.H.Kim & C.H.Kim—Ophioglossaceae. Based on the diagnosis associated with Mankyua B.Y.Sun, M.H.Kim & C.H.Kim, Taxon 50(4): 1020, f. 1. 2002. Phegopteridoideae Salino, A.R.Sm. & T.E.Almeida, subfam.
e (1852)—Thelypteridaceae. nov. Type: Phegopteris (C.Presl) Fe Laminae bipinnate-pinnatifid or more divided, or laminae bipinnatifid or tripinnatifid; costae adaxially lacking grooves; veins free, not reaching the margins; chromosome base numbers x ¼ 30 or 31. Pteridineae J.Prado & Schuettp., stat. nov.  Pteridaceae E.D. M.Kirchn., Schul-Bot. 109. 1831. Type: Pteris L. (1753)— Polypodiales. Saccolomatineae Hovenkamp, subord. nov. Type: Saccoloma Kaulf. (1820)—Saccolomataceae. Rhizome erect, covered with multistratose, non-clathrate scales, in cross-section with two concentric rings of meristeles, sori cup-shaped, spores trilete, tetrahedral. Acknowledgements We thank all pteridologists, past and present, who have provided the taxonomic and phylogenetic foundation for this effort. We are also grateful to Kanchi Gandhi for helping with some nomenclatural aspects. www.jse.ac.cn PPG I References Adjie B, Lestari WS. 2014. A phylogenetic placement of dicksonioid fern Calochlaena javanica (Dicksoniaceae). Floribunda 5: 17–20. Adjie B, Takamiya M, Ohta M, Ohsawa TA, Watano Y. 2008. Molecular phylogeny of the lady fern genus Athyrium in Japan based on chloroplast rbcL and trnL-trnF sequences. Acta Phytotaxonomica et Geobotanica 59: 79–95. Almeida TE, Hennequin S, Schneider H, Smith AR, Batista JAN, Ramalho AJ, Proite K, Salino A. 2016. Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688–700. APG. 1998. An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85: 531–553. 595 Chao YS, Rouhan G, Amoroso VB, Chiou W-L. 2014. Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae). Annals of Botany 114: 109–124. Chase MW, Reveal JL. 2009. A phylogenetic classification of the land plants to accompany APG III. Botanical Journal of the Linnean Society 161: 122–127. Chen CW, Schuettpelz E, Lindsay S, Middleton DJ. 2016. Proposal to conserve the name Haplopteris against Monogramma (Pteridaceae). Taxon 65: 884–885. Christenhusz MJM. 2007. Dracoglossum, a new neotropical fern genus (Pteridophyta). Thaiszia 17: 1–10. Christenhusz MJM. 2009. Marattiidae (23 January 2009) [online]. In: Tree of Life Web Project, available from http://tolweb.org/ Marattiales/21664/2009.01.23 [accessed 7 October 2016]. Christenhusz MJM. 2012. Pteridaceae. In: Greuter W, von RaabStraube E eds. EuroþMed Notulae, 6. Willdenowia 42: 284. APG II. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: 399–436. Christenhusz MJM, Chase MW. 2014. Trends and concepts in fern classification. Annals of Botany 113: 571–594. APG III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. APG III. Botanical Journal of the Linnean Society 161: 105–121. Christenhusz MJM, Jones M, Lehtonen S. 2013. Phylogenetic placement of the enigmatic fern genus Dracoglossum. American Fern Journal 103: 131–138. APG IV. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20. Christenhusz MJM, Schneider H. 2011. Corrections to Phytotaxa 19: linear sequence of lycophytes and ferns. Phytotaxa 28: 50–52. Assis FC, Almeida TE, Russell SJ, Schneider H, Salino A. 2016. Molecular phylogeny and recircumscription of the fern genus Pecluma (Polypodiaceae-Polypodiopsida). Phytotaxa 247: 235–246. Christenhusz MJM, Tuomisto H, Metzgar JS, Pryer KM. 2008. Evolutionary relationships within the neotropical, eusporangiate fern genus Danaea (Marattiaceae). Molecular Phylogenetics and Evolution 46: 34–48. Backlund A, Bremer K. 1998. To be or not to be: Principles of classification and monotypic plant families. Taxon 47: 391–400. Beck JB, Windham MD, Yatskievych G, Pryer KM. 2010. A diploids-first approach to species delimitation and interpreting polyploid evolution in the fern genus Astrolepis (Pteridaceae). Systematic Botany 35: 223–234. Benham DM, Windham MD. 1992. Generic affinities of the star-scaled cloak ferns. American Fern Journal 82: 47–58. Bernhardi JJ. 1806. Dritter Versuch einer Anordnung der Farrnkr€ auter. Neues Journal f€ ur die Botanik 1 (2): 1–50. Bishop LE. 1977. The American species of Grammitis sect. Grammitis. American Fern Journal 67: 101–106. Bishop LE. 1978. Revision of the genus Cochlidium (Grammitidaceae). American Fern Journal 68: 76–94. Bishop LE. 1988. Ceradenia, a new genus of Grammitidaceae. American Fern Journal 78: 1–5. Bishop LE. 1989. Zygophlebia, a new genus of Grammitidaceae. American Fern Journal 79: 103–118. Bishop LE, Smith AR. 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New World. Systematic Botany 117: 345–362. Bomfleur B, Grimm GW, McLoughlin S. 2015. Osmunda pulchella sp. nov. from the Jurassic of Sweden—reconciling molecular and fossil evidence in the phylogeny of modern royal ferns (Osmundaceae). BMC Evolutionary Biology 15: 126. Christenhusz MJM, Zhang X-C, Schneider H. 2011. A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. Christensen C. 1916. Maxonia, a new genus of tropical American ferns. Smithsonian Miscellaneous Collections 66: 1–4. Christensen C, Ching RC. 1934. Pteridrys, a new fern genus from tropical Asia. Bulletin of the Fan Memorial Institute of Biology; Botany 5: 125–145. Clausen RT. 1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19: 1–177. Cochran AT, Prado J, Schuettpelz E. 2014. Tryonia, a new taenitidoid fern genus segregated from Jamesonia and Eriosorus (Pteridaceae). PhytoKeys 35: 23–43. Conant DS. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. Journal of the Arnold Arboretum 64: 333–382. Copeland EB. 1947. Genera filicum, the genera of ferns. Annales Cryptogamici et Phytopathologici 5: 1–247 [Genera Filicum]. Cranfill R, Kato M. 2003. Phylogenetics, biogeography, and classification of the woodwardioid ferns (Blechnaceae). In: Pteridology in the New Millennium. Springer Netherlands: 25–48. Dauphin B, Vieu J, Grant JR. 2014. Molecular phylogenetics supports widespread cryptic species in moonworts (Botrychium s.s., Ophioglossaceae). American Journal of Botany 101: 128–140. Brownsey PJ. 1983. Polyploidy and aneuploidy in Hypolepis and the evolution of the Dennstaedtiales. American Fern Journal 73: 97–108. Der JP, Thomson JA, Stratford JK, Wolf PG. 2009. Global chloroplast phylogeny and biogeography of bracken (Pteridium; Dennstaedtiaceae). American Journal of Botany 96: 1041–1049. Cantino PD, de Queiroz K. 2000. PhyloCode: A phylogenetic code of biological nomenclature [online]. Available from https://www. ohio.edu/PhyloCode/PhyloCode.pdf [accessed 7 October 2016]. Des Marais DL, Smith AR, Britton DM, Pryer KM. 2003. Phylogenetic relationships and evolution of extant horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F). International Journal of Plant Sciences 164: 737–751. C
ardenas GG, Tuomisto H, Lehtonen S. 2016. Newly discovered diversity in the tropical fern genus Metaxya based on morphology and molecular phylogenetic analyses. Kew Bulletin 71: 5–31. www.jse.ac.cn Du X-Y, Cheng X. 2011. Phylogeny of the Chinese endemic fern genus Neocheiropteris (Polypodiaceae). Plant Diversity 33: 261–268. J. Syst. Evol. 54 (6): 563–603, 2016 596 The Pteridophyte Phylogeny Group Duan Y-F, Hennequin S, Rouhan G, Bassuner B, Zhang L-B. 2017. A taxonomic revision of the fern genus Ctenitis (Dryopteridaceae) from Africa and the western Indian Ocean. Annals of the Missouri Botanical Garden 102: in press. Dubuisson J-Y, Hennequin S, Douzery EJ, Cranfill RB, Smith AR, Pryer KM. 2003. rbcL phylogeny of the fern genus Trichomanes (Hymenophyllaceae), with special reference to neotropical taxa. International Journal of Plant Sciences 164: 753–761. Ebihara A. 2011. rbcL phylogeny of Japanese pteridophyte flora and implications on infrafamilial systematics. Bulletin of the National Museum of Nature and Science, Series B 37: 63–74. Ebihara A, Dubuisson J-Y, Iwatsuki K, Hennequin S, Ito M. 2006. A taxonomic revision of Hymenophyllaceae. Blumea 51: 221–280. Ebihara A, Iwatsuki K, Ito M, Hennequin S, Dubuisson J-Y. 2007. A global molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linnean Society 155: 1–27. Ebihara A, Nakato N, Amoroso VB, Hidayat A, Kuo L-Y. 2016. Monachosorum arakii Tagawa (Dennstaedtiaceae) is a relict “international” hybrid: A reassessment of the Monachosorum species. Systematic Botany 41: 586–595. Eiserhardt WL, Rohwer JG, Russell SJ, Yesilyurt JC, Schneider H. 2011. Evidence for radiations of cheilanthoid ferns in the Greater Cape Floristic Region. Taxon 60: 1269–1283. Field AR, Bostock PD. 2013. New and existing combinations in Palaeotropical Phlegmariurus (Lycopodiaceae) and lectotypification of the type species Phlegmariurus phlegmaria (L.) T. Sen & U. Sen. PhytoKeys 20: 33–51. Hall CC, Lellinger DB. 1967. A revision of the fern genus Mildella. American Fern Journal 57: 113–134. Haufler CH, Pryer KM, Schuettpelz E, Sessa EB, Farrar DR, Moran R, Schneller JJ, Watkins Jr JE, Windham MD. 2016. Sex and the single gametophyte: Revising the homosporous vascular plant life cycle in light of contemporary research. BioScience. doi: 10.1093/biosci/ biw108. Hauk WD, Parks CR, Chase MW. 2003. Phylogenetic studies of Ophioglossaceae: Evidence from rbcL and trnL-F plastid DNA sequences and morphology. Molecular Phylogenetics and Evolution 28: 131–151. Hauke RL. 1990. Equisetaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 46–48. Hayata B. 1927. On the systematic importance of the stelar system in the Filicales, I. Botanical Magazine, Tokyo 41: 697–718. Hayata B. 1928. On the systematic importance of the stelar system in the Filicales, III. Botanical Magazine, Tokyo 42: 334–348. He H, Wu S, Xiang J, Barrington DS. 2013. Arachniodes. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 542–558. He L-J, Zhang X-C. 2012. Exploring generic delimitation within the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution 65: 757–764. He Z, Kato M. 2013. Diplaziopsidaceae. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 317–318. Field AR, Testo W, Bostock PDB, Holtum JAM, Waycott M. 2016. Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum. Molecular Phylogenetics and Evolution 94: 635–657. Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson J-Y. 2006. New insights into the phylogeny of the genus Hymenophyllum s.l. (Hymenophyllaceae): Revealing the polyphyly of Mecodium. Systematic Botany 31: 271–284. Gasper AL, Almeida TE, Dittrich VAO, Smith AR, Salino A. 2016a. Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus-level treatment. Cladistics. doi: 10.1111/ cla.12173. Hennequin S, Hovenkamp P, Christenhusz MJM, Schneider H. 2010. Phylogenetics and biogeography of Nephrolepis—a tale of old settlers and young tramps. Botanical Journal of the Linnean Society 164: 113–127. Gasper AL, Dittrich VAO, Smith AR, Salino A. 2016b. A classification for Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Hennequin S, Kessler M, Lindsay S, Schneider H. 2014. Evolutionary patterns in the assembly of fern diversity on the oceanic Mascarene Islands. Journal of Biogeography 41: 1651–1663. Gastony GJ, Ungerer MC. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae). American Journal of Botany 84: 840–849. Hennipman E, Veldhoen P, Kramer KU, Price MG. 1990. Polypodiaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 203–230. Geiger JM, Korall P, Ranker TA, Kleist AC, Nelson CL. 2013. Molecular phylogenetic relationships of Cibotium and origin of the Hawaiian endemics. American Fern Journal 103: 141–152. Greuter W, Troia A. 2014. Proposal to conserve the name Palhinhaea against Lepidotis. Taxon 63: 680–681. Grusz AL, Windham MD. 2013. Toward a monophyletic Cheilanthes: The resurrection and recircumscription of Myriopteris (Pteridaceae). PhytoKeys 32: 49–64. Hirai RY, Rouhan G, Labiak PH, Ranker TA, Prado J. 2011. Moranopteris: A new neotropical genus of grammitid ferns (Polypodiaceae) segregated from Asian Micropolypodium. Taxon 60: 1123–1137. Holttum RE, Grimes JW. 1980. The genus Pseudocyclosorus Ching (Thelypteridaceae). Kew Bulletin 34: 499–516. Holttum RE. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the Old World. Blumea 19: 17–52. Grusz AL, Windham MD, Yatskievych G, Huiet L, Gastony GJ, Pryer KM. 2014. Patterns of diversification in the xeric-adapted fern genus Myriopteris (Pteridaceae). Systematic Botany 39: 698– 714. Holttum RE. 1974a. Thelypteridaceae of Africa and adjacent islands. Journal of South African Botany 40: 123–168. Guillon JM. 2007. Molecular phylogeny of horsetails (Equisetum) including chloroplast atpB sequences. Journal of Plant Research 120: 569–574. Holttum RE. 1975. Studies in the family Thelypteridaceae VIII. The genera Mesophlebion and Plesioneuron. Blumea 22: 223–250. Haines A. 2003. The families Huperziaceae and Lycopodiaceae in New England: A taxonomic and ecological reference. Bar Harbor: V.F. Thomas Co. J. Syst. Evol. 54 (6): 563–603, 2016 Holttum RE. 1974b. The fern-genus Pleocnemia. Kew Bulletin 29: 341–357. Holttum RE. 1986. Studies in the fern-genera allied to Tectaria Cav. VI. A conspectus of genera in the Old World regarded as related to Tectaria, with descriptions of two genera. The Gardens’ Bulletin Singapore 39: 153–167. www.jse.ac.cn PPG I Holub J. 1964. Lycopodiella, novy rod radu Lycopodiales. Preslia 36: 16–22. Holub J. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 97–110. Holub J. 1983. Validation of generic names in Lycopodiaceae: With a description of a new genus Pseudolycopodiella. Folia Geobotanica Phytotaxonomica 18: 439–442. Holub J. 1985. Transfers of Lycopodium species to Huperzia: With a note on generic classification in Huperziaceae. Folia Geobotanica Phytotaxonomica 20: 67–80. Holub J. 1991. Some taxonomic changes within the Lycopodiales. Folia Geobotanica Phytotaxonomica 26: 81–94. Hoot SB, Taylor WC, Napier NS. 2006. Phylogeny and biogeography of Isoëtes (Isoëtaceae) based on nuclear and chloroplast DNA sequence data. Systematic Botany 31: 449–460. H€ orandl E, Stuessy TF. 2010. Paraphyletic groups as natural units of biological classification. Taxon 59: 1641–1653. Hovenkamp PH, Miyamoto F. 2012. Nephrolepidaceae. In: Nooteboom HP ed. Flora Malesiana Series (Series II—Ferns and fern allies). Leiden: Flora Malesiana Foundation. 4: 97–122. Hovenkamp PH. 1997. Paraselliguea, a new genus of Malesian Polypodiaceae. Blumea 42: 485–487. Humphreys AE, Linder HP. 2009. Concept versus data in delimitation of plant genera. Taxon 58: 1054–1074. Jermy AC. 1990a. Isoëtaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 26–31. Jermy AC. 1990b. Selaginellaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 39–45. Jessen S, Lehmann L, Bujnock W. 2012. Cryptogramma bithynica spec. nov. (Pteridaceae, Pteridophyta)—A new fern species from Northwestern Anatolia/Turkey. Fern Gazette 19 (2): 47–54. Ji SG, Huo KK, Wang J, Pan SL. 2008. A molecular phylogenetic study of Huperziaceae based on chloroplast rbcL and psbAtrnH sequences. Journal of Systematics and Evolution 46: 213–219. 597 Korall P, Kenrick P. 2004. The phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: Extreme substitution rates and rate heterogeneity. Molecular Phylogenetics and Evolution 31: 852–864. Korall P, Kenrick P, Therrien JP. 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. International Journal of Plant Sciences 160: 585–594. Korall P, Pryer KM, Metzgar JS, Schneider H, Conant DS. 2006. Tree ferns: Monophyletic groups and their relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics and Evolution 39: 830–845. Kramer KU. 1990a. Psilotaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 22–25. Kramer KU. 1990b. Cheiropleuriaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 68–69. Kramer KU. 1990c. Davalliaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 74–80. Kramer KU. 1990d. Dennstaedtiaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 81–94. Kramer KU. 1990e. Dicksoniaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 94–99. Kramer KU. 1990f. Dipteridaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 99–101. Kramer KU. 1990g. Gleicheniaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 145–152. Kramer KU. 1990h. Lomariopsidaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 164–170. Johnson AK, Rothfels CJ, Windham MD, Pryer KM. 2012. Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae). American Journal of Botany 99: 1118–1124. Kramer KU. 1990i. Lophosoriaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 170–172. Kato M, Setoguchi H. 1998. An rbcL-based phylogeny and heteroblastic leaf morphology of Matoniaceae. Systematic Botany 23: 391–400. Kramer KU. 1990j. Loxomataceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 172–174. Kato M, Yatabe Y, Sahashi N, Murakami N. 2001. Taxonomic studies of Cheiropleuria (Dipteridaceae). Blumea 46: 513–525. Kramer KU. 1990k. Marsileaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 180–183. Kenrick P, Crane PR. 1997. The origin and early diversification of land plants: A cladistic study. Washington: Smithsonian. Kessler M, Vel
asquez ALM, Sundue M, Labiak PH. 2011. Alansmia, a new genus of grammitid ferns (Polypodiaceae) segregated from Terpsichore. Brittonia 63: 233–244. Knie N, Fischer S, Grewe F, Polsakiewicz M, Knoop V. 2015. Horsetails are the sister group to all other monilophytes and Marattiales are sister to leptosporangiate ferns. Molecular Phylogenetics and Evolution 90: 140–149. Koenemann DM, Maisonpierre JA, Barrington DS. 2011. Broad-scale integrity and local divergence in the fiddlehead fern Matteuccia struthiopteris (L.) Todaro (Onocleaceae). American Fern Journal 101: 213–230. Korall P, Conant DS, Metzgar JS, Schneider H, Pryer KM. 2007. A molecular phylogeny of scaly tree ferns (Cyatheaceae). American Journal of Botany 94: 873–886. www.jse.ac.cn Kramer KU. 1990l. Matoniaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 183–185. Kramer KU. 1990m. Monachosoraceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 187–188. Kramer KU. 1990n. Oleandraceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 190–193. Kramer KU. 1990o. Osmundaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 197–200. Kramer KU. 1990p. Plagiogyriaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 201–203. J. Syst. Evol. 54 (6): 563–603, 2016 598 The Pteridophyte Phylogeny Group Kramer KU. 1990q. Schizaeaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 258–263. Kramer KU, Green PS eds. 1990. Pteridophytes and gymnosperms. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I: 1–404. Kramer KU, Holttum RE, Moran RC, Smith AR. 1990. Dryopteridaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 101–144. Kreier HP, Rex M, Weising K, Kessler M, Smith AR, Schneider H. 2008a. Inferring the diversification of the epiphytic fern genus Serpocaulon (Polypodiaceae) in South America using chloroplast sequences and amplified fragment length polymorphisms. Plant Systematics and Evolution 274: 1–6. Kreier HP, Rojas-Alvarado AF, Smith AR, Schneider H. 2007. Hyalotrichopteris is indeed a Campyloneurum (Polypodiaceae). American Fern Journal 97: 127–135. Kreier HP, Schneider H. 2006a. Reinstatement of Loxogramme dictyopteris, based on phylogenetic evidence, for the New Zealand endemic fern, Anarthropteris lanceolata (Polypodiaceae, Polypodiidae). Australian Systematic Botany 19: 309–314. Kreier HP, Schneider H. 2006b. Phylogeny and biogeography of the staghorn fern genus Platycerium (Polypodiaceae, Polypodiidae). American Journal of Botany 93: 217–225. Kreier HP, Zhang XC, Muth H, Schneider H. 2008b. The microsoroid ferns: Inferring the relationships of a highly diverse lineage of Paleotropical epiphytic ferns (Polypodiaceae, Polypodiopsida). Molecular Phylogenetics and Evolution 48: 1155–1167. Kuo L-Y, Chang Y-H, Glowienka JM, Amoroso VB, Dong S-Y, Kao T-T, Wang C-N, Chiou W-L. 2016a. A revised framework of Dryopteris subg. Nothoperanema (Dryopteridaceae) inferred from phylogenetic evidence, with descriptions of two new sections. Systematic Botany 41: 596–605. Kuo L-Y, Ebihara A, Shinohara W, Rouhan G, Wood KR, Wang C-N, Chiou W-L. 2016b. Historical biogeography of the fern genus Deparia (Athyriaceae) and its relation with polyploidy. Molecular Phylogenetics and Evolution 104: 123–134. Kuo L-Y, Li F-W, Chiou W-L, Wang C-N. 2011. First insights into fern matK phylogeny. Molecular Phylogenetics and Evolution 59: 556–566. Labiak PH. 2011. Stenogrammitis, a new genus of grammitid ferns segregated from Lellingeria (Polypodiaceae). Brittonia 63: 139–149. Le P
echon T, He H, Zhang L, Zhou X-M, Gao X-F, Zhang L-B. 2016. Using a multilocus phylogeny to test morphology-based classifications of Polystichum (Dryopteridaceae), one of the largest fern genera. BMC Evolutionary Biology 16: 55. Lehnert M, M€ onnich M, Pleines T, Schmidt-Lebuhn A, Kessler M. 2001. The relictual fern genus Loxsomopsis. American Fern Journal 91: 13–24. Lehnert M. 2013. Grammitid ferns (Polypodiaceae): Melpomene, Flora Neotropica Monograph, 112. New York: The New York Botanical Garden. Lehtonen S. 2011. Towards resolving the complete fern tree of life. PLoS One 6: e24851. Lehtonen S, Tuomisto H, Rouhan G, Christenhusz MJM. 2010. Phylogenetics and classification of the pantropical fern family Lindsaeaceae. Botanical Journal of the Linnean Society 163: 305–359. Lehtonen S, Tuomisto H, Rouhan G, Christenhusz MJM. 2013. Taxonomic revision of the fern genus Osmolindsaea (Lindsaeaceae). Systematic Botany 38: 887–900. Lehtonen S, Wahlberg N, Christenhusz MJM. 2012. Diversification of lindsaeoid ferns and phylogenetic uncertainty of early polypod relationships. Botanical Journal of the Linnean Society 170: 489–503. Le
on B. 1995. Campyloneurum. In: Moran RC, Riba R eds. Flora Mesoamericana. Mexico City: Universidad Aut
onoma de M
exico. I (Psilotaceae to Salviniaceae): 333–338. Li C, Lu S, Ma J, Sun X, Gai Y, Barrington DS, Yang Q. 2012a. From the Himalayan region or the Malay Archipelago: Molecular dating to trace the origin of a fern genus Phymatopteris (Polypodiaceae). Chinese Science Bulletin 57: 4569–4577. Li C, Lu S, Ma J, Yang Q. 2010. Phylogeny and divergence of Gleicheniaceae inferred from three plastid genes. Acta Palaeontologica Sinica 49: 64–72. Li C, Lu S, Sun X, Yang Q. 2011. Phylogenetic positions of the enigmatic asiatic fern genera Diplaziopsis and Rhachidosorus from analyses of four plastid genes. American Fern Journal 101: 142–155. Li F-W, Pryer KM, Windham MD. 2012b. Gaga, a new fern genus segregated from Cheilanthes (Pteridaceae). Systematic Botany 37: 845–860. Li F-W, Tan BC, Buchbender B, Moran RC, Rouhan G, Wang CN, Quandt D. 2009. Identifying a mysterious aquatic fern gametophyte. Plant Systematics and Evolution 281: 77–86. Labiak PH. 2013. Grammitid ferns (Polypodiaceae): Lellingeria, Flora Neotropica Monograph 111. New York: The New York Botanical Garden. Lin Y, Li Z, Iwatsuki K, Smith AR. 2013. Thelypteridaceae. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 319–396. Labiak PH, Mickel JT, Hanks JG. 2015a. Molecular phylogeny and character evolution of Anemiaceae (Schizaeales). Taxon 64: 1141–1158. Lindsay S, Middleton DJ, Suksathan P. 2012. A new species of Rhachidosorus (Rhachidosoraceae), a genus new to Thailand. Thai Forest Bulletin (Botany) 40: 102–104. Labiak PH, Rouhan G, Sundue M. 2010. Phylogeny and taxonomy of Leucotrichum (Polypodiaceae): A new genus of grammitid ferns from the Neotropics. Taxon 59: 911–921. Link-P
erez MA, Hickey RJ. 2011. Revision of Adiantopsis radiata (Pteridaceae) with descriptions of new taxa with palmately compound laminae. Systematic Botany 36: 565–582. Labiak PH, Sundue M, Rouhan G, Hanks JG, Mickel JT, Moran RC. 2014. Phylogeny and historical biogeography of the lastreopsid ferns (Dryopteridaceae). American Journal of Botany 101: 1207–1228. Link-P
erez MA, Watson LE, Hickey RJ. 2011. Redefinition of Adiantopsis F
ee (Pteridaceae): Systematics, diversification, and biogeography. Taxon 60: 1255–1268. Labiak PH, Sundue M, Rouhan G, Moran RC. 2015b. New combinations in Lastreopsis and Parapolystichum (Dryopteridaceae). Brittonia 67: 79–86. Liu H-M, Zhang X-C, Wang M-P, Shang H, Zhou S-L, Yan Y-H, Wei X-P, Xu W-B, Schneider H. 2016. Phylogenetic placement of the enigmatic fern genus Trichoneuron informs on the infra-familial relationship of Dryopteridaceae. Plant Systematics and Evolution 302: 319–332. Lars
en E, Rydin C. 2016. Disentangling the phylogeny of Isoetes (Isoetales), using nuclear and plastid data. International Journal of Plant Sciences 177: 157–174. J. Syst. Evol. 54 (6): 563–603, 2016 Liu H, Jiang R-H, Guo J, Hovenkamp P, Perrie LR, Shepherd L, Hennequin S, Schneider H. 2013. Towards a phylogenetic www.jse.ac.cn PPG I 599 classification of the climbing fern genus Arthropteris. Taxon 62: 688–700. Murillo MT, Smith AR. 2003. Luisma, a new genus of Grammitidaceae (Pteridophyta) from Colombia. Novon 13: 313–317. L
origa J, Schmidt AR, Moran RC, Feldberg K, Schneider H, Heinrichs J. 2014. The first fossil of a bolbitidoid fern belongs to the earlydivergent lineages of Elaphoglossum (Dryopteridaceae). American Journal of Botany 101: 1466–1475. Mynssen CM, Vasco A, Moran RC, Sylvestre LS, Rouhan G. 2016. Desmophlebiaceae and Desmophlebium: A new family and genus of eupolypod II ferns. Taxon 65: 19–34. Lu J-M, Barrington DS, Li D-Z. 2007. Molecular phylogeny of the polystichoid ferns in Asia based on rbcL sequences. Systematic Botany 32: 26–33. Nagalingum NS, Nowak MD, Pryer KM. 2008. Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia. Botanical Journal of the Linnean Society 157: 673–685. McKeown M, Sundue M, Barrington D. 2012. Phylogenetic analyses place the Australian monotypic Revwattsia in Dryopteris (Dryopteridaceae). PhytoKeys 14: 43. Nagalingum NS, Schneider H, Pryer KM. 2007. Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus Marsilea. Systematic Botany 32: 16–25. McNeill J, Pryer KM. 1985. The status and typification of Phegopteris and Gymnocarpium. Taxon 34: 136–143. Nakazato T, Gastony GJ. 2003. Molecular phylogenetics of Anogramma species and related genera (Pteridaceae: Taenitidoideae). Systematic Botany 28: 490–502. Metzgar JS, Alverson ER, Chen S, Vaganov AV, Ickert-Bond SM. 2013. Diversification and reticulation in the circumboreal fern genus Cryptogramma. Molecular Phylogenetics and Evolution 67: 589–599. Metzgar JS, Schneider H, Pryer KM. 2007. Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae). International Journal of Plant Sciences 168: 1045–1053. Metzgar JS, Skog JE, Zimmer EA, Pryer KM. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany 33: 31–36. Mickel JT. 2016. Anemia (Anemiaceae), Flora Neotropica Monograph, 118. New York: The New York Botanical Garden. Mickel JT, Smith AR. 2004. The Pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: 1–1054. Miller Jr CN. 1967. Evolution of the fern genus Osmunda. Contributions from the Museum of Paleontology, University of Michigan 21: 139–203. Moran RC. 1986. The neotropical fern genus Olfersia. American Fern Journal 76: 161–178. Moran RC. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural History Survey Bulletin 34: 1–138. Moran RC. 1991. Monograph of the neotropical fern genus Stigmatopteris (Dryopteridaceae). Annals of the Missouri Botanical Garden 78: 857–914. Moran RC. 2000. Monograph of the neotropical species of Lomariopsis (Lomariopsidaceae). Brittonia 52: 55–111. Moran RC, Labiak PH. 2015. Phylogeny of the polybotryoid fern clade (Dryopteridaceae). International Journal of Plant Sciences 176: 880–891. Moran RC, Labiak PH. 2016. Phylogeny and character evolution of the neotropical fern genus Stigmatopteris (Dryopteridaceae). Brittonia. doi: 10.1007/s12228-016-9437-1. Moran RC, Labiak PH, Hanks JG, Prado J. 2014. The phylogenetic relationship of Tectaria brauniana and Tectaria nicotianifolia, and the recognition of Hypoderris (Tectariaceae). Systematic Botany 39: 384–395. Øllgaard B. 1990. Lycopodiaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 31–39. Øllgaard B. 2012a. Nomenclatural changes in Brazilian Lycopodiaceae. Rodrigu
esia 63: 479–482. Øllgaard B. 2012b. New combinations in neotropical Lycopodiaceae. Phytotaxa 57: 10–22. Øllgaard B. 2015. Six new species and some nomenclatural changes in neotropical Lycopodiaceae. Nordic Journal of Botany 33: 186–196. Øllgaard B, Windisch PG. 2014. Lycopodiaceae in Brazil. Conspectus of the family I. The genera Lycopodium, Austrolycopodium, Diphasium, and Diphasiastrum. Rodrigu
esia 65: 261–277. Otto EM, Janßen T, Kreier HP, Schneider H. 2009. New insights into the phylogeny of Pleopeltis and related neotropical genera (Polypodiaceae, Polypodiopsida). Molecular Phylogenetics and Evolution 53: 190–201. Pab
on-Mora N, Gonz
alez F. 2015. Nephopteris out of the clouds: molecular evidence places the enigmatic N. maxonii (Pteridaceae) within the Jamesonia clade. Brittonia 68: 83–92. Parris BS. 1990. Grammitidaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 153–157. Parris BS. 1997. Themelium, a new genus of Grammitidaceae. Kew Bulletin 52: 737–741. Parris BS. 1998. Chrysogrammitis, a new genus of Grammitidaceae (Filicales). Kew Bulletin 53: 909–918. Parris BS. 2004. New combinations in Acrosorus, Lellingeria, Prosaptia and Themelium (Grammitidaceae: Filicales). Kew Bulletin 59: 223–225. Parris BS. 2007. Five new genera and three new species of Grammitidaceae (Filicales) and the re-establishment of Oreogrammitis. The Gardens’ Bulletin Singapore 58: 233–274. Parris BS. 2010a. Prosaptia. In: Flora of Peninsular Malaysia, ser. 1. 1: 170–182. Moran RC, Labiak PH, Sundue M. 2010a. Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae). International Journal of Plant Sciences 171: 547–559. Parris BS. 2010b. New combinations in Oreogrammitis, Prosaptia, Radiogrammitis, Themelium and Tomophyllum (Grammitidaceae: Polypodiopsida: Monilophyta) from Malesia and the Pacific Islands. Kew Bulletin 65: 123–125. Moran RC, Labiak PH, Sundue M. 2010b. Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia 62: 337–356. Parris BS. 2013. Archigrammitis, a new genus of grammitid fern (Polypodiaceae) from Malesia and Polynesia. Fern Gazette 19: 135–138. Murdock AG. 2008. A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new genus Ptisana. Taxon 57: 737–755. Perrie LR, Bayly MJ, Lehnebach CA, Brownsey PJ. 2007. Molecular phylogenetic and molecular dating of the New Zealand Gleicheniaceae. Brittonia 59: 129–141. www.jse.ac.cn J. Syst. Evol. 54 (6): 563–603, 2016 600 The Pteridophyte Phylogeny Group Perrie LR, Brownsey PJ, Lovis JD. 2010. Tmesipteris horomaka, a new octoploid species from Banks Peninsula. New Zealand Journal of Botany 48: 15–29. Rakotondrainibe F. 2010. Le genre Rumohra Raddi (Pteridophyta, Dryopteridaceae) a Madagascar: une esp ece et deux combinaisons nouvelles. Adansonia 32: 217–228. Perrie LR, Parris BS. 2012. Chloroplast DNA sequences indicate the grammitid ferns (Polypodiaceae) in New Zealand belong to a single clade, Notogrammitis gen. nov. New Zealand Journal of Botany 50: 457–472. Ramos-Giacosa JP, Morbelli MA, Giudice GE. 2011. Spore morphology and wall ultrastructure of Trachypteris species (Pteridaceae). Plant Systematics and Evolution 294: 227–237. Perrie LR, Shepherd LD, Brownsey PJ. 2015. An expanded phylogeny of the Dennstaedtiaceae ferns: Oenotrichia falls within a nonmonophyletic Dennstaedtia, and Saccoloma is polyphyletic. Australian Systematic Botany 28: 256–264. Ranker TA. 2008. A new combination in Adenophorus (Polypodiaceae). American Fern Journal 98: 170–177. Ranker TA, Haufler CH. 1990. A new combination in Bommeria (Adiantaceae). American Fern Journal 80: 1–3. Perrie LR, Wilson RK, Shepherd LD, Ohlsen DJ, Batty EL, Brownsey PJ, Bayly MJ. 2014. Molecular phylogenetics and generic taxonomy of Blechnaceae ferns. Taxon 63: 745–758. Ranker TA, Smith AR, Parris BS, Geiger JM, Haufler CH, Straub SC, Schneider H. 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): A case of rampant morphological homoplasy. Taxon 53: 415–428. Pichi Sermolli REG. 1973. Historical review of the higher classification of the Filicopsida. In: Jermy AC, Crabbe JA, Thomas BA eds. The phylogeny and classification of the ferns. Botanical Journal of the Linnean Society 67 (supplement 1): 11–40. Reid JD, Plunkett GM, Peters GA. 2006. Phylogenetic relationships in the heterosporous fern genus Azolla (Azollaceae) based on DNA sequence data from three noncoding regions. International Journal of Plant Sciences 167: 529–538. Pichi Sermolli REG. 1977a. Fragmenta Pteridologiae—VI. Webbia 31: 246–247. Rothfels CJ, Larsson A, Kuo L-Y, Korall P, Chiou W-L, Pryer KM. 2012a. Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Systematic Biology 61: 490–509. Pichi Sermolli REG. 1977b. Tentamen pteridophytorum genera in taxanomicum ordinam redigendi. Webbia 31: 315–512. Prado J, Del Nero R, Salatino A, Salatino ML. 2007. Phylogenetic relationships among Pteridaceae, including Brazilian species, inferred from rbcL sequences. Taxon 56: 355–368. Prado J, Moran RC. 2008. Revision of the neotropical species of Triplophyllum (Tectariaceae). Brittonia 60: 103–130. Prado J, Moran RC. 2016. Monograph of the West Indian fern genus Polystichopsis (Dryopteridaceae). Brittonia 68: 1–24. Pryer KM. 1999. Phylogeny of marsileaceous ferns and relationships of the fossil Hydropteris pinnata reconsidered. International Journal of Plant Sciences 160: 931–954. Pryer KM, Huiet L, Li F-W, Rothfels CJ, Schuettpelz E. 2016. Maidenhair ferns, Adiantum, are indeed monophyletic and sister to shoestring ferns, vittarioids (Pteridaceae). Systematic Botany 41: 17–23. Rothfels CJ, Li F-W, Sigel EM, Huiet L, Larsson A, Burge DO, Ruhsam M, Deyholos M, Soltis DE, Stewart Jr. CN, Shaw SW, Pokorny L, Chen T, dePamphilis C, DeGironimo L, Chen L, Wei X, Sun X, Korall P, Stevenson DW, Graham SW, Wong GK-S, Pryer KM. 2015. The evolutionary history of ferns inferred from 25 low-copy nuclear genes. American Journal of Botany 102: 1–19. Rothfels CJ, Schuettpelz E. 2014. Accelerated rate of molecular evolution for vittarioid ferns is strong and not driven by selection. Systematic Biology 63: 31–54. Rothfels CJ, Sundue MA, Kuo L-Y, Larsson A, Kato M, Schuettpelz E, Pryer KM. 2012b. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61: 515–533. Pryer KM, Schneider H, Smith AR, Cranfill R, Wolf PG, Hunt JS. 2001. Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Nature 409: 618–621. Rothfels CJ, Windham MD, Grusz AL, Gastony GJ, Pryer KM. 2008. Toward a monophyletic Notholaena (Pteridaceae): Resolving patterns of evolutionary convergence in xeric-adapted ferns. Taxon 57: 712–724. Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R. 2004. Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. American Journal of Botany 91: 1582–1598. Rothfels CJ, Windham MD, Pryer KM. 2013. A plastid phylogeny of the cosmopolitan fern family Cystopteridaceae (Polypodiopsida). Systematic Botany 38: 295–306. Pryer KM, Smith AR, Rothfels C. 2008. Polypodiidae (23 December 2008) [online]. In: Tree of Life Web Project, available from http://tolweb.org/Polypodiidae/21666/2008.12.23 [accessed 7 October 2016]. Pryer KM, Smith AR, Rothfels C. 2009. Polypodiopsida (14 January 2009) [online]. In: Tree of Life Web Project: http://tolweb.org/ Polypodiopsida/20615/2009.01.14 [accessed 7 October 2016]. Pryer KM, Smith AR, Skog JE. 1995. Phylogenetic relationships of extant ferns based on evidence from morphology and rbcL sequences. American Fern Journal 85: 205–282. Qiu YL, Li L, Wang B, Chen Z, Dombrovska O, Lee J, Kent L, Li R, Jobson RW, Hendry TA, Taylor DW. 2007. A nonflowering land plant phylogeny inferred from nucleotide sequences of seven chloroplast, mitochondrial, and nuclear genes. International Journal of Plant Sciences 168: 691–708. Rai HS, Graham SW. 2010. Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes). American Journal of Botany 96: 1444–1456. J. Syst. Evol. 54 (6): 563–603, 2016 Rouhan G, Dubuisson J-Y, Rakotondrainibe F, Motley TJ, Mickel JT, Labat J-N, Moran RC. 2004. Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: Contributions of species from the Indian Ocean area. Molecular Phylogenetics and Evolution 33: 745–763. Rouhan G, Hanks JG, McClelland D, Moran RC. 2007. Preliminary phylogenetic analysis of the fern genus Lomariopsis (Lomariopsidaceae). Brittonia 59: 115–128. Rouhan G, Labiak PH, Randrianjohany E, Rakotondrainibe F. 2012. Not so neotropical after all: The grammitid fern genus Leucotrichum (Polypodiaceae) is also paleotropical, as revealed by a new species from Madagascar. Systematic Botany 37: 331–338. Roux JP. 2001. Conspectus of Southern African Pteridophyta (Southern African Botanical Diversity Network Report, 3). Pretoria: Sabonet. Ruggiero MA, Gordon DP, Orrell TM, Bailly N, Bourgoin T, Brusca RC, Cavalier-Smith T, Guiry MD, Kirk PM. 2015. A higher level classification of all living organisms. PLoS ONE 10: e0119248. www.jse.ac.cn PPG I Ruhfel BR, Gitzendanner MA, Soltis PS, Soltis DE, Burleigh JG. 2014. From algae to angiosperms–inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes. BMC Evolutionary Biology 14: 23. Rydin C, Wikstr€ om N. 2002. Phylogeny of Isoëtes (Lycopsida): Resolving basal relationships using rbcL sequences. Taxon 51: 83–89. Salino A, Almeida TE, Smith AR, Gomez AN, Kreier HP, Schneider H. 2008. A new species of Microgramma (Polypodiaceae) from Brazil and recircumscription of the genus based on phylogenetic evidence. Systematic Botany 33: 630–635. 601 Schuettpelz E, Pryer KM, Windham MD. 2015. A unified approach to taxonomic delimitation in the fern genus Pentagramma (Pteridaceae). Systematic Botany 40: 629–644. Schuettpelz E, Pryer KM. 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037–1050. Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM. 2007. A molecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera. Molecular Phylogenetics and Evolution 44: 1172–1185. Sarvela J. 1978. A synopsis of the fern genus Gymnocarpium. Annales Botanici Fennici 15: 101–106. Schwartsburd PB, Prado J. 2015. A taxonomic revision of the South American species of Hypolepis (Dennstaedtiaceae), Part I. American Fern Journal 105: 263–313. Schmidt-Lebuhn AN. 2011. Fallacies and false premises—a critical assessment of the arguments for the recognition of paraphyletic taxa in botany. Cladistics 28: 174–187. Schwartsburd PB, Prado J. 2016. A taxonomic revision of the South American species of Hypolepis (Dennstaedtiaceae), Part II. American Fern Journal 106: 1–53. Schneider H, He L, Hennequin S, Zhang X-C. 2013. Towards a natural classification of Pteridaceae: Inferring the relationships of enigmatic pteridoid fern species occurring in the Sino-Himalaya and Afro-Madagascar. Phytotaxa 77: 49–60. Sessa EB, Zimmer EA, Givnish TJ. 2012. Phylogeny, divergence times, and historical biogeography of New World Dryopteris (Dryopteridaceae). American Journal of Botany 99: 730–750. Schneider H, Kreier HP, Hovenkamp P, Janssen T. 2008. Phylogenetic relationships of the fern genus Christiopteris shed new light onto the classification and biogeography of drynarioid ferns. Botanical Journal of the Linnean Society 157: 645–656. Schneider H, Kreier HP, Wilson E, Smith AR. 2006. The Synammia enigma: Evidence for a temperate lineage of polygrammoid ferns (Polypodiaceae, Polypodiidae) in southern South America. Systematic Botany 31: 31–41. Schneider H, Ranker TA, Russell SJ, Cranfill R, Geiger JM, Aguraiuja R, Wood KR, Grundmann M, Kloberdanz K, Vogel JC. 2005. Origin of the endemic fern genus Diellia coincides with the renewal of Hawaiian terrestrial life in the Miocene. Proceedings of the Royal Society B: Biological Sciences 272: 455–460. Schneider H, Russell SJ, Cox CJ, Bakker F, Henderson S, Gibby M, Vogel JC. 2004a. Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its implications for the biogeography. Systematic Botany 29: 260–274. Schneider H, Schuettpelz E, Pryer KM, Cranfill R, Magall
on S, Lupia R. 2004b. Ferns diversified in the shadow of angiosperms. Nature 428: 553–557. Schneider H, Smith AR, Cranfill R, Hildebrand T, Haufler CH, Ranker T. 2004c. Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): Exploring aspects of the diversification of epiphytic plants. Molecular Phylogenetics and Evolution 31: 1041–1063. Schneider H, Smith AR, Pryer KM. 2009. Is morphology really at odds with molecules in estimating fern phylogeny? Systematic Botany 34: 455–475. Shao Y, Wei R, Zhang X, Xiang Q. 2015. Molecular phylogeny of the cliff ferns (Woodsiaceae: Polypodiales) with a proposed infrageneric classification. PLoS ONE 10: e0136318. Shinohara W, Nakato N, Yatabe-Kakugawa Y, Oka T, Kim JK, Murakami N, Noda H, Sahashi N. 2013. The use of matK in Ophioglossaceae phylogeny and the determination of Mankyua chromosome number shed light on chromosome number evolution in Ophioglossaceae. Systematic Botany 38: 564–570. Shmakov AI. 2015. The new system of family Woodsiaceae. Turczaninowia 18: 11–16. Sigel EM, Windham MD, Haufler CH, Pryer KM. 2014. Phylogeny, divergence time estimates, and phylogeography of the diploid species of the Polypodium vulgare complex (Polypodiaceae). Systematic Botany 39: 1042–1055. Sigel EM, Windham MD, Huiet L, Yatskievych G, Pryer KM. 2011. Species relationships and farina evolution in the cheilanthoid fern genus Argyrochosma (Pteridaceae). Systematic Botany 36: 554–564. Smith AR. 1986. Revision of the neotropical fern genus Cyclodium. American Fern Journal 76: 56–98. Smith AR. 1990. Thelypteridaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 263–272. Smith AR. 1992. A review of the fern genus Micropolypodium (Grammitidaceae). Novon 2: 419–425. Smith AR. 1993. Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3: 478–489. Smith AR. 1995. Non-molecular phylogenetic hypotheses for ferns. American Fern Journal 85: 104–122. Schneller JJ. 1990a. Azollaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 57–60. Smith AR, Cranfill RB. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae). American Fern Journal 92: 131–149. Schneller JJ. 1990b. Salviniaceae. In: Kubitzki K ed. The Families and Genera of Vascular Plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 256–258. Smith AR, Kreier HP, Haufler CH, Ranker T, Schneider H. 2006a. Serpocaulon (Polypodiaceae), a new genus segregated from Polypodium. Taxon 55: 919–930. Schuettpelz E, Chen CW, Kessler M, Pinson JB, Johnson G, Davila A, Cochran AT, Huiet L, Pryer KM. 2016. A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data. Taxon 65: 708–722. Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG. 2006b. A classification for extant ferns. Taxon 55: 705–731. Schuettpelz E, Davila A, Prado J, Hirai RY, Yatskievych G. 2014. Molecular phylogenetic and morphological affinities of Adiantum senae (Pteridaceae). Taxon 63: 258–264. www.jse.ac.cn Smith AR, Tejero-Diez D. 2014. Pleopeltis (Polypodiaceae), a redefinition of the genus and nomenclatural novelties. Botanical Sciences 92: 43–58. Stuessy TF. 2009. Plant taxonomy: The systematic evaluation of comparative data. New York: Columbia University Press. J. Syst. Evol. 54 (6): 563–603, 2016 602 The Pteridophyte Phylogeny Group Sun B-Y, Baek TG, Kim Y-D, Kim CS. 2009. Phylogeny of the family Ophioglossaceae with special emphasis on genus Mankyua. Korean Journal of Plant Taxonomy 39: 135–142. Sundue MA. 2010. A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae). Brittonia 62: 357–399. Sundue MA. 2013. Mycopteris, a new neotropical genus of grammitid ferns (Polypodiaceae). Brittonia 66: 174–185. Sundue MA, Hirai RY, Prado J. 2013. Rumohra glandulosissima (Dryopteridaceae) a new species from the Atlantic Rainforest, and revision of the species occurring in Brazil. Systematic Botany 38: 915–924. Sundue MA, Labiak PH, Mostacero J, Smith AR. 2012. Galactodenia, a new genus of grammitid ferns segregated from Terpsichore (Polypodiaceae). Systematic Botany 37: 339–246. Wei R, Schneider H, Zhang X-C. 2013. Toward a new circumscription of the twinsorus-fern genus Diplazium (Athyriaceae): A molecular phylogeny with morphological implications and infrageneric taxonomy. Taxon 62: 441–457. Wei R, Xiang Q-P, Schneider H, Sundue MA, Kessler M, Kamau PW, Hidayat A, Zhang X-C. 2015. Eurasian origin, boreotropical migration and transoceanic dispersal in the pantropical fern genus Diplazium (Athyriaceae). Journal of Biogeography 42: 1809–1819. Wei R, Zhang X-C, Qi X-P. 2010. Phylogeny of Diplaziopsis and Homalosorus based on two chloroplast DNA sequences: rbcL and rps4þrps4-trnS IGS. Acta Botanica Yunnanica 17: 46–54. Wei R, Zhang X-C. 2014. Rediscovery of Cystoathyrium chinense Ching (Cystopteridaceae): Phylogenetic placement of the critically endangered fern species endemic to China. Journal of Systematics and Evolution 52: 450–457. Sundue MA, Parris BS, Ranker TA, Smith AR, Fujimoto EL, ZamoraCrosby D, Morden CW, Chiou W-L, Chen C-W, Rouhan G, Hirai RY, Prado J. 2014. Global phylogeny and biogeography of grammitid ferns (Polypodiaceae). Molecular Phylogenetics and Evolution 81: 195–206. Weststrand S, Korall P. 2016a. Phylogeny of Selaginellaceae: There is value in morphology after all! American Journal of Botany. doi: 10.3732/ajb.1600156. Sundue M, Testo W. 2016. Parapolystichum novoguineense (comb. nov; Dryopteridaceae) from New Guinea. Phytotaxa 243: 193– 196. Weststrand S, Korall P. 2016b. A subgeneric classification of Selaginella (Selaginellaceae). American Journal of Botany. doi: 10.3732/ ajb.1600288. Thomson JA. 2012. Taxonomic status of diploid southern hemisphere brackens (Pteridium: Dennstaedtiaceae). Telopea 14: 43–48. White RA, Turner MD. 1988. Calochlaena, a new genus of dicksonioid ferns. American Fern Journal 78: 86–95. Tindale MD. 1965. A monograph of the genus Lastreopsis Ching. Contributions of the New South Wales National Herbarium 3: 249–339. Whitten WM, Jacono CC, Nagalingum NS. 2012. An expanded plastid phylogeny of Marsilea with emphasis on North American species. American Fern Journal 102: 114–135. Tryon RM. 1970. The classification of the Cyatheaceae. Contributions from the Gray Herbarium of Harvard University 200: 3–53. Wickett NJ, Mirarab S, Nguyen N, Warnow T, Carpenter E, Matasci N, Ayyampalayam S, Barker MS, Burleigh JG, Gitzendanner MA, Ruhfel BR. 2014. Phylotranscriptomic analysis of the origin and early diversification of land plants. Proceedings of the National Academy of Sciences USA 111: E4859–4868. Tryon RM, Tryon AF, Kramer KU. 1990. Pteridaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 230–256. Wikstr€ om N, Kenrick P. 1997. Phylogeny of Lycopodiaceae (Lycopsida) and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. International Journal of Plant Sciences 158: 862–871. Tsutsumi C, Chen C-W, Larsson A, Hirayama Y, Kato M. 2016. Phylogeny and classification of Davalliaceae based on chloroplast and nuclear markers. Taxon 65: in press Wikstr€ om N, Kenrick P. 2000. Relationships of Lycopodium and Lycopodiella based on combined plastid rbcL gene and trnL intron sequence data. Systematic Botany 25: 495–510. Tsutsumi C, Kato M. 2006. Evolution of epiphytes in Davalliaceae and related ferns. Botanical Journal of the Linnean Society 151: 495–510. Wikstr€ om N, Kenrick P. 2001. Evolution of Lycopodiaceae (Lycopsida): Estimating divergence times from rbcL gene sequences by use of nonparametric rate smoothing. Molecular Phylogenetics and Evolution 19: 177–186. Tryon RM. 1962. Taxonomic fern notes. III. Contributions from the Gray Herbarium of Harvard University 191: 91–107. Wagner Jr WH. 1990. Ophioglossaceae. In: Kubitzki K ed. The families and genera of vascular plants. Berlin: Springer-Verlag. I (Kramer KU, Green PS eds. Pteridophytes and gymnosperms): 193–197. Wagner Jr WH, Beitel JM. 1992. Generic classification of modern North American Lycopodiaceae. Annals of the Missouri Botanical Garden 79: 676–686. Wagner Jr WH, Wagner FS. 1980. Polyploidy in pteridophytes. In: Lewis WH ed. Polyploidy. New York: Springer. 199–214. Wang F-G, Barratt S, Falc
on W, Fay MF, Lehtonen S, Tuomisto H, Xing F-W, Christenhusz MJM. 2014. On the monophyly of subfamily Tectarioideae (Polypodiaceae) and the phylogenetic placement of some associated fern genera. Phytotaxa 164: 1–16. Wang L, Wu ZQ, Xiang QP, Heinrichs J, Schneider H, Zhang XC. 2010. A molecular phylogeny and a revised classification of tribe Lepisoreae (Polypodiaceae) based on an analysis of four plastid DNA regions. Botanical Journal of the Linnean Society 162: 28–38. Wang W, Yang W, Mao X, Zhao R, Dou P, Zhang G. 2015. The phylogenetic affinities of Pellaea connectens, a rare endemic Chinese fern. Phytotaxa 220: 30–42. J. Syst. Evol. 54 (6): 563–603, 2016 Wikstr€ om N, Kenrick P, Vogel JC. 2002. Schizaeaceae: A phylogenetic approach. Review of Palaeobotany and Palynology 119: 35–50. Windham MD. 1987. Argyrochosma, a new genus of cheilanthoid ferns. American Fern Journal 77: 37–41. Wolf PG. 1995. Phylogenetic analyses of rbcL and nuclear ribosomal RNA gene sequences in Dennstaedtiaceae. American Fern Journal 85: 306–327. Wolf PG, Rowe CA, Der JP, Schilling MP, Visger CJ, Thomson JA. 2015. Origins and diversity of a cosmopolitan fern genus on an island archipelago. AoB Plants 7: plv118. Wood TE, Takebayashi N, Barker MS, Mayrose I, Greenspoon PB, Rieseberg LH. 2009. The frequency of polyploid speciation in vascular plants. Proceedings of the National Academy of Sciences USA 106: 13875–13879. Yatabe Y, Murakami N, Iwatsuki K. 2005. Claytosmunda: A new subgenus of Osmunda (Osmundaceae). Acta Phytotaxonomica et Geobotanica 56: 127–128. www.jse.ac.cn PPG I Yatabe Y, Nishida H, Murakami N. 1999. Phylogeny of Osmundaceae inferred from rbcL nucleotide sequences and comparison to the fossil evidences. Journal of Plant Research 112: 397–404. Yatskievych G, Arbelaez AL. 2008. A new species and three generic transfers in the fern genus Notholaena (Pteridaceae). Novon 18: 120–124. Yatskievych G, Smith AR. 2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52: 331–336. Yatskievych G, Stein DB, Gastony GJ. 1988. Chloroplast DNA evolution and systematics of Phanerophlebia (Dryopteridaceae) and related fern genera. Proceedings of the National Academy of Sciences USA 85: 2589–2593. Yesilyurt JC, Barbara T, Schneider H, Russell S, Culham A, Gibby M. 2015. Identifying the generic limits of the cheilanthoid genus Doryopteris (Pteridaceae). Phytotaxa 221: 101–122. Yesilyurt JC, Schneider H. 2010. The new fern genus Calciphilopteris (Pteridaceae). Phytotaxa 7: 52–59. Zhang G, Ranker TA. 2013a. Parahemionitis. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 235. Zhang G, Ranker TA. 2013b. Paragymnopteris. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 235–237. Zhang G, Yatskievych G. 2013. Cheilanthes. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 218–224. Zhang G, Yatskievych G, Hooper EA. 2013a. Aleuritopteris. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 224–235. Zhang L, Rothfels CJ, Ebihara A, Schuettpelz E, Le P
echon T, Kamau P, He H, Zhou X-M, Prado J, Field A, Yatskievych G, Gao X-F, Zhang LB. 2015. A global plastid phylogeny of the brake fern genus Pteris (Pteridaceae) and related genera in the Pteridoideae. Cladistics 31: 406–423. Zhang L, Schuettpelz E, Rothfels CJ, Zhou X-M, Gao X-F, Zhang L-B. 2016. Circumscription and phylogeny of the fern family Tectariaceae based on plastid and nuclear markers, with the description of two new genera: Draconopteris and Malaifilix (Tectariaceae). Taxon 65: 723–738. www.jse.ac.cn 603 Zhang L-B, Iwatsuki K. 2013. Lycopodiaceae. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 13–34. Zhang L-B, Kung H-S. 1998. A taxonomic study of Huperzia Berhn. (s.s.) sect. Huperzia in China. Acta Phytotaxonomica Sinica 36: 521–529. Zhang L-B, Kung H-S. 1999. On the taxonomy of Phlegmariurus (Herter) Holub (Huperziaceae) sect. Huperzioides H.S. Kung et L.B. Zhang (sect. nov.) in China. Acta Phytotaxonomica Sinica 37: 40–53. Zhang L-B, Kung H-S. 2000a. Taxonomy of Huperzia Bernh. (sen. str.) sect. Serratae (Rothm.) Holub in China. Acta Phytotaxonomica Sinica 38: 13–22. Zhang L-B, Kung H-S. 2000b. Two sections of Phlegmariurus (Herter) Holub (Huperziaceae) from China. Acta Phytotaxonomica Sinica 38: 23–29. Zhang L-B, Wu S, Xiang J, Xing F, He H, Wang F, Lu S, Dong S, Barrington DS, Iwatsuki K, Christenhusz MJM, Mickel JT, Kato M, Gilbert MG. 2013b. Dryopteridaceae. In: Wu Z, Raven PH, Hong D eds. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. 2–3 (Lycopodiaceae through Polypodiaceae): 541–724. Zhang L-B, Zhang L. 2012. The inclusion of Acrophorus, Diacalpe, Nothoperanema, and Peranema in Dryopteris: The molecular phylogeny, systematics, and nomenclature of Dryopteris subg. Nothoperanema (Dryopteridaceae). Taxon 61: 1199–1216. Zhang L-B, Zhang L. 2015. Didymochlaenaceae: A new fern family of eupolypods I (Polypodiales). Taxon 64: 27–38. Zhang L-B, Zhang L, Dong S-Y, Sessa EB, Gao X-F, Ebihara A. 2012. Molecular circumscription and major evolutionary lineages of the fern genus Dryopteris (Dryopteridaceae). BMC Evolutionary Biology 12: 180. Zhou S, Dong W, Chen X, Zhang X, Wen J, Schneider H. 2014. How many species of bracken (Pteridium) are there? Assessing the Chinese brackens using molecular evidence. Taxon 63: 509–521. Zhou XM, Rothfels CJ, Zhang L, He ZR, Le P
echon T, He H, Lu NT, Knapp R, Lorence D, He XJ, Gao XF, Zhang L-B. 2015. A large-scale phylogeny of the lycophyte genus Selaginella (Selaginellaceae: Lycopodiopsida) based on plastid and nuclear loci. Cladistics 32: 360–389. Zhou XM, Zhang LB. 2015. A classification of Selaginella (Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features. Taxon 64: 1117–1140. J. Syst. Evol. 54 (6): 563–603, 2016