Aclassification for extant ferns - Sites@Duke · leptosporangiate ferns, evidence is now sufficient...

INTRODUCTIONRecent phylogenetic studies have revealed a basal

dichotomy within vascular plants separating the lyco-phytes (less than 1 of extant vascular plants) from theeuphyllophytes (Fig 1 Raubeson amp Jansen 1992 Ken-rick amp Crane 1997 Pryer amp al 2001a 2004a b) Liv-ing euphyllophytes in turn comprise two major cladesthe spermatophytes (seed plants) which are in excess of260000 species (Thorne 2002 Scotland amp Wortley2003) and the monilophytes (ferns sensu Pryer amp al2004b) with about 9000 species including horsetailswhisk ferns and all eusporangiate and leptosporangiateferns Plants that are included in the lycophyte and fernclades are all spore-bearing or ldquoseed-freerdquo and becauseof this common feature their members have been lumpedtogether historically under various terms such as ldquopteri-dophytesrdquo and ldquoferns and fern alliesrdquomdashparaphyleticassemblages of plants The focus of this reclassificationis exclusively on ferns (Division Tracheophyta Sub-division Euphyllophytina Infradivision Moniliformop-ses of Kenrick amp Crane 1997) characterized by lateralroot origin in the endodermis usually mesarch pro-toxylem in shoots a pseudoendospore plasmodial tape-tum and sperm cells with 30ndash1000 flagellae (Renzagliaamp al 2000 Schneider amp al 2002a)

Increasingly robust phylogenetic hypotheses for

ferns (Hasebe amp al 1994 1995 Manhart 1994 1995Pryer amp al 1995 2001a 2004b Kranz amp Huss 1996Pahnke amp al 1996 Wolf 1997 Wolf amp al 1998Beckert amp al 1999 Vangerow amp al 1999 Sano amp al2000a Schneider amp al 2004c Wikstroumlm amp Pryer 2005Tsutsumi amp Kato 2006 Schuettpelz amp al in press) uti-

705

Smith amp al bull Fern classification55 (3) bull August 2006 705ndash731

Fig 1 Consensus phylogeny depicting relationships ofmajor vascular plant lineages Topology summarizes theresults of previously published phylogenetic studies(eg Raubeson amp Jansen 1992 Kenrick amp Crane 1997Renzaglia amp al 2000 Pryer amp al 2001a see main text forothers) Resolved nodes have received bootstrap supportge 70

Euphyllophytes

MonilophytesSpermatophytesLycophytes

TAXONOMY

A classification for extant ferns

Alan R Smith1 Kathleen M Pryer2 Eric Schuettpelz2 Petra Korall23 Harald Schneider4 ampPaul G Wolf5

1 University Herbarium 1001 Valley Life Sciences Building 2465 University of California BerkeleyCalifornia 94720-2465 USA arsmithberkeleyedu (author for correspondence)

2 Department of Biology Duke University Durham North Carolina 27708-0338 USA3 Department of Phanerogamic Botany Swedish Museum of Natural History Box 50007 SE-104 05 Stock-

holm Sweden4 Albrecht-von-Haller-Institut fuumlr Pflanzenwissenschaften Abteilung Systematische Botanik Georg-August-

Universitaumlt Untere Karspuumlle 2 37073 Goumlttingen Germany5 Department of Biology Utah State University Logan Utah 84322-5305 USA

We present a revised classification for extant ferns with emphasis on ordinal and familial ranks and a synop-sis of included genera Our classification reflects recently published phylogenetic hypotheses based on bothmorphological and molecular data Within our new classification we recognize four monophyletic classes 11monophyletic orders and 37 families 32 of which are strongly supported as monophyletic One new familyCibotiaceae Korall is described The phylogenetic affinities of a few genera in the order Polypodiales areunclear and their familial placements are therefore tentative Alphabetical lists of accepted genera (includingcommon synonyms) families orders and taxa of higher rank are provided

KEYWORDS classification Cibotiaceae ferns monilophytes monophyletic

lizing data from morphology seven chloroplast markers(rbcL atpA atpB accD rps4 16S rDNA ITS) one nu-clear gene (18S rDNA) and three mitochondrial genes(atp1 nad2 nad5) prompt us to reevaluate the classifi-cation of these vascular plants Multiple-gene phyloge-netic analyses eg studies by Wolf (1996) Wolf amp al(1998) Pryer amp al (2001a 2004b) Schneider amp al(2004c) Wikstroumlm amp Pryer (2005) and Schuettpelz amp al(in press) have given rise to growing conviction in boththe composition and relationships of taxa at familial andordinal ranks Five recent morphological analyses ofrelationships by Pryer amp al (1995 2001a) Schneider(1996a) Stevenson amp Loconte (1996) and Schneider ampal (in prep) have increased support for the molecular-based consensus topology For eusporangiate and basalleptosporangiate ferns evidence is now sufficient to al-low us to circumscribe confidently most clades and as-sign ranks However for some more derived leptospo-rangiate ferns the phylogenetic evidence is still some-what equivocal Consequently declaration of phyloge-netic positions for some taxa and the assignment of ranks(which we consider subjective and secondary) in a clas-sical ldquoLinnaean-stylerdquo hierarchy are tentative In this pa-per we present a revised view of the classification of ex-tant ferns taking into account all relevant evidence Wefocus our classification at the ranks of class order andfamily believing that the information at hand is mostappropriate for resolution and understanding of relation-ships at these levels Within most families and especial-ly at the generic level there is still insufficient evidenceto attempt many classificatory decisions

In the classification proposed herein we account forand place in a revised taxonomic framework all names atfamily and ordinal rank utilized in previous major classifi-cations directed at ferns particularly those that have beenproposed in the last eighty years Bower (1926) Christen-sen (1938) Ching (1940 1978) Dickason (1946) Cope-land (1947) Holttum (1947 1949 1973) Pichi Sermolli(1958 1977) Mehra (1961) Wagner (1969) Nayar(1970) Tagawa amp Iwatsuki (1972) Mickel (1974) Try-on amp Tryon (1982) Kramer (in Kubitzki 1990) Hennip-man (1996) and Stevenson amp Loconte (1996) We alsoconsider various herbarium schemes in common usesuch as the one by Crabbe amp al (1975) Many of theseclassifications as well as others have been reviewed andreproduced by Pichi Sermolli (1973) who also summa-rized detailed information on family names of ferns(1970 1981 1982 1986) Smith (1995) provided a re-cent summary and discussion of these classifications

Classifications serve many purposes among them toprovide a genealogical framework in which to identifyplants organize herbaria retrieve information and toconduct many kinds of studies (eg evolutionary mor-phological and physiological) Multi-use classifications

serve us best if we name only those clades that are read-ily recognizable and characterized by morphologicalsynapomorphic characters at least at family and higherranks However more traditional (morphology-based)and practical classifications are sometimes incompatiblewith the results and classifications implied by phyloge-netic studies especially when the principle of monophy-ly is used as a grouping criterion (recognizing clades andnot paraphyletic grades APG II 2003) When a tradi-tionally recognized family nests within another complexclassificatory choices ensue (1) recognition of para-phyletic families (Brummitt 1996 1997 Lideacuten amp al1997 Moore 1998 Diggs amp Lipscomb 2002 Grant2003) (2) dismemberment of a recognized family intosmaller families (eg the disintegration of classicalScrophulariaceae Olmstead amp al 2001) or (3) integra-tion of the traditional family that causes the paraphylyinto the ldquoprogenitorrdquo family The first choice preferredby some leads to recognition of unnatural (non-mono-phyletic) groups which in our opinion often retards orobscures investigation into interesting biological phyto-geographic and evolutionary questions The second so-lution supposes that we have morphological synapomor-phies for nodes that lead to all of the segregate familiesand this is not often the case although one hopes thateventually we shall find these synapomorphies Untilthen it may be nearly impossible to define some segre-gate families in such a way that they would be bothkeyable and circumscribable The third solution is a ldquofastfixrdquo to the problem but expediency often demands thatat least some of the intrafamilial subclades also be rec-ognized taxonomically either at a lower rank (eg sub-family) or with an unranked informal name (eg ldquogram-mitid fernsrdquo) until further decisions on rank can be madeThis third option seems to us the most practicable andpractical solution toward a ldquofirst-passrdquo revision of fernclassification As more data are gathered and future phy-logenetic analyses provide better resolved and better sup-ported topologies one expects further insight into identi-fying synapomorphies for segregate taxa enabling even-tual movement toward the second option of recognizingsegregate monophyletic families

The nested linearity of a ranked Linnaean classifica-tion and the availability of a limited number of univer-sally accepted ranks facilitate the organization and ar-rangement of taxa for cataloging purposes such as theefficient arrangement of specimens in herbaria or theorganization and retrieval of regional biodiversity fromchecklists floras and other taxonomic products Rankedclassifications often perform poorly however whencalled upon to present explicit statements of clade mem-bership and relationship Such classifications also canmislead one into making specious comparisons eg at-tempting to describe the extent of biodiversity in terms of

Smith amp al bull Fern classification 55 (3) bull August 2006 705ndash731

706

numbers of taxa of a particular arbitrarily designatedrank (comparing apples with oranges) Such problemsnotwithstanding it is nevertheless highly likely thatranked Linnaean classifications will continue to co-existwith phylogenetic classifications for the foreseeablefuture

In this paper we combine the principle of monophy-ly with a desire to maintain well-established names toupdate ordinal and familial ranks within ferns so thatthey are better reconciled with our current best estimatesof phylogenetic relationships However we attempt toavoid recognizing monogeneric families within the morederived leptosporangiate ferns To paraphrase Crisp amp al(1999) who addressed the matter of monotypic generathe most compelling case for recognizing a monotypicfamily is when such a family is characterized by one ormore autapomorphies and is the sister group to a clade offamilies all of the members of which lack these apomor-phies The alternative would be to treat the whole clade(including the autapomorphic family) as a single familywhich results in sinking some well-circumscribed fami-lies into synonymy At this time we prefer to utilize aminimum number of ranks to categorize only the mostwell supported splits in the phylogeny

In some respects our new classification differs inrelatively minor ways in the circumscription of ordersfamilies and hierarchical structure from most other re-cent fern classifications (eg those of Tryon amp Tryon1982 Kramer in Kubitzki 1990 and Stevenson ampLoconte 1996) but there are significant departures Onlyone previous fern classification has employed cladisticmethodology in a rigorous way Stevenson amp Loconte(1996) superimposed on their tree a hierarchical classifi-cation (albeit somewhat abbreviated) but the phylogenythat they generated was based exclusively on morpho-logical data and differs radically from the most up-to-date phylogenetic hypothesis Our classification in con-trast is based on consensus of a variety of morphologi-cal and molecular studies

Fern names above the rank of genus used in this clas-sification (Appendices 1ndash4) have been obtained for themost part from the web site of James Reveal Universityof Maryland (httpwwwlifeumdeduemeritusrevealPBIOfamhightaxaindexhtml) and from Hoogland ampReveal (2005) Most names at family rank are also listedand discussed by Pichi Sermolli (1970 1982) and sum-marized in reports by the Subcommittee for FamilyNames of Pteridophyta (Pichi Sermolli 1981 1986) thislist was further emended and updated by Pichi Sermolli(1993) To the best of our knowledge (and unless other-wise indicated) the family names in Appendix 3 have allbeen published in accord with the International Code ofBotanical Nomenclature (Greuter amp al 2000) and thussatisfy the relevant Articles in the Code with regard to

publication and priority of family names (Art 106 Art111ndash113 Art 18 Art 411) and names of ranks abovefamily (Art 16 17) Although the principle of priority isnot mandatory for names of taxa above the rank of fam-ily (Art 119) authors are advised to follow this princi-ple (Rec 16B1) Of the orders we recognize below allhave priority except Salviniales which is later than themuch less commonly applied name Pilulariales

The rules of nomenclature applicable to supra-ordi-nal names are the same as those for ordinal names dis-cussed above We have made no attempt to scrutinize theliterature for names above the rank of order but many ofthese names are included in lists available from theReveal website (and so are included in Appendix 1) andin Hoogland amp Reveal (2005) still others can be gleanedfrom Kenrick amp Crane (1997 see especially their Table72 pp 231ndash233 however many of these are not valid-ly published) and from Pichi Sermolli (1959) Names atordinal and subordinal ranks also mostly from the Re-veal website and from Hoogland amp Reveal (2005) aregiven in Appendix 2 names at familial rank are given inAppendix 3 Citations for all names are given in Ap-pendix 4 We also present an index to commonly accept-ed genera with family assignments proposed here(Appendix 5)

In the classification that follows for each family wegive common names (often derived from the scientificnames) for the clades (if there is one that has commonusage) heterotypic synonyms approximate numbers ofgenera and species names of constituent genera refer-ences to relevant phylogenetic literature and discussionof unresolved problems where appropriate DNA se-quence data are now available for all families recognizedherein and for most genera of ferns A superscript num-ber one (1) denotes those genera for which DNA se-quence data are not available nonetheless taxonomicplacement for most of these is relatively certain based onmorphological evidence Lack of a superscript indicatesthat some molecular evidence (either published orunpublished) has been available to us for considerationand we are relatively confident in the placement of thegenus The classification presented below is based on theconsensus relationships depicted in Fig 2 which arederived from and guided by recent and ongoing phylo-genetic studies (eg Hasebe amp al 1995 Pryer amp al2004b Schneider amp al 2004d Korall amp al 2006Schuettpelz amp al 2006 unpubl data)

CLASSIFICATION OF EXTANTFERNS

In this reclassification we treat all classes ordersand families of extant ferns which constitute a mono-


707

phyletic group sometimes referred to as InfradivisionMoniliformopses (Kenrick amp Crane 1997) or monilo-phytes (Donoghue in Judd amp al 2002 Pryer amp al2001a 2004a b) However ldquoInfradivisionrdquo is not a rec-

ognized rank in the International Code of Botanical No-menclature (Greuter amp al 2000) moreover the nameldquoMoniliformopsesrdquo was never validly published lackinga Latin diagnosis or description or a reference to one


Fig 2 Consensus phylogeny depicting fern relationships upon which our classification is based Topology summar-izes the results of previously published and ongoing phylogenetic studies (eg Hasebe amp al 1995 Pryer amp al 2004bSchneider amp al 2004b Korall amp al 2006 Schuettpelz amp al in press Schuettpelz unpubl see main text for others)Resolved nodes have received bootstrap support ge 70 except those drawn with dotted lines Family order and classnames that correspond to our classification are indicated to the right Common names for some larger clades are indi-cated to the left

PsilotaceaeOphioglossaceae

PsilotopsidaPsilotalesOphioglossales

EquisetopsidaEquisetalesEquisetaceaeMarattiopsidaMarattialesMarattiaceae

Polypodio-psida

OsmundalesOsmundaceae

GleichenialesGleicheniaceaeDipteridaceaeMatoniaceae

HymenophyllalesHymenophyllaceae

SchizaealesSchizaeaceae

LygodiaceaeAnemiaceae

SalvinialesSalviniaceaeMarsileaceae

CyathealesPlagiogyriaceae

LoxomataceaeThyrsopteridaceae

CibotiaceaeCyatheaceae

Polypodiales

LindsaeaceaeSaccolomataceaeDennstaedtiaceaePteridaceaeAspleniaceae

ThelypteridaceaeWoodsiaceae

OnocleaceaeBlechnaceae

Dryopteridaceae

DavalliaceaePolypodiaceae

Leptosporangiates

Core leptosporangiates

Ferns

Polypods

Eupolypods

Eupolypods II

Eupolypods I

MetaxyaceaeDicksoniaceae

Culcitaceae

OleandraceaeTectariaceaeLomariopsidaceae

708

Because validly published names at an appropriate rankare not available for ferns (as here defined) euphyllo-phytes (ferns + seed plants) or even vascular plants (eu-phyllophytes + lycophytes) and because all availablenames at higher ranks have been used in a sense we thinkwould be confusing we avoid placing ferns in any rankabove class Classification may be better served by adop-ting ldquofloatingrdquo rankless names for clades above the rankof class

Within ferns we recognize four classes (Psilotop-sida Equisetopsida Marattiopsida Polypodiopsida) 11orders and 37 families

I CLASS PSILOTOPSIDAA ORDER OPHIOGLOSSALES1 Family Ophioglossaceae mdash Ophioglosso-

ids incl Botrychiaceae HelminthostachyaceaeFour genera Botrychium (grapeferns moonworts) Hel-minthostachys Mankyua1 Ophioglossum (addertongues) Botrychium (incl Botrychium ss Sceptridi-um Botrypus and Japanobotrychium) and Ophioglos-sum (incl Cheiroglossa Ophioderma) are sometimes di-vided more finely (Kato 1987 Hauk amp al 2003) Ca 80spp monophyletic (Hasebe amp al 1995 Hauk 1995Pryer amp al 2001a 2004b Hauk amp al 2003) Mankyuafrom Cheju Island Korea has recently been describedbut no molecular data are available (Sun amp al 2001)Species mostly terrestrial (a few epiphytic) temperateand boreal but a few pantropical Characters vernationnodding (not circinate) rhizomes and petioles fleshyroot hairs lacking aerophores absent (Davies 1991) fer-tile leaves each with a single sporophore arising at thebase of or along the trophophore stalk or at the base ofthe trophophore blade (several sporophores per blade inCheiroglossa) sporangia large with walls two cellsthick lacking an annulus spores globose-tetrahedral tri-lete many (gt 1000) per sporangium gametophytes sub-terranean non-photosynthetic mycorrhizal x = 45 (46)

B ORDER PSILOTALES2 Family Psilotaceae mdash Whisk ferns incl

Tmesipteridaceae Two genera (Psilotum Tmesipte-ris) ca 12 total spp (2 in Psilotum) monophyletic (Ha-sebe amp al 1995 Pryer amp al 2001a 2004) Charactersroots absent stems bearing reduced unveined or single-veined euphylls sporangia large with walls two cellsthick lacking an annulus two or three sporangia fused toform a synangium seemingly borne on the adaxial sideof a forked leaf spores reniform monolete many (gt1000) per sporangium gametophytes subterranean (Psi-lotum) non-photosynthetic mycorrhizal x = 52

II CLASS EQUISETOPSIDA [= SPHE-NOPSIDA]

C ORDER EQUISETALES3 Family Equisetaceae mdash Horsetails A single

genus (Equisetum) 15 spp usually placed in two well-marked subgenera subg Equisetum and subg Hippo-chaete monophyletic (Pryer amp al 2001a 2004b DesMarais amp al 2003 Guillon 2004) The spermatozoidsof Equisetum share several important features with otherferns that support their inclusion in this clade (Renzagliaamp al 2000) Kato (1983) adduced additional morpho-logical characters including root characters supporting arelationship between horsetails and ferns Charactersstems whorled lacunate leaves whorled connate spo-rangia with helical secondary wall thickenings (Bate-man 1991) borne on peltate sporangiophores that col-lectively comprise strobili sporangia large lacking anannulus many (gt 1000) per sporangium spores greenwith circular aperture and four paddle-like coiled ela-ters gametophytes green surficial x = 108

III CLASS MARATTIOPSIDAD ORDER MARATTIALES mdash Including

Christenseniales4 Family Marattiaceae mdash Marattioids incl

Angiopteridaceae Christenseniaceae DanaeaceaeKaulfussiaceae Four genera Angiopteris Christen-senia Danaea Marattia Archangiopteris has been rec-ognized by some (eg Pichi Sermolli 1977) but appearsto nest within Angiopteris (Murdock 2005) Danaea issister to the other three genera (Pryer amp al 2001a2004b Murdock 2005) and represents a neotropicalradiation (Christenhusz amp al unpubl) Angiopteris andChristensenia are restricted to eastern and southeasternAsia Australasia and Polynesia while Marattia is pan-tropical Ca 150 spp but monographic revision is need-ed at the species level in several genera monophyletic(Hill amp Camus 1986 Pryer amp al 2001a 2004bMurdock 2005) We see no advantage or good reason forrecognizing several of the constituent genera as mono-generic families as done by Pichi Sermolli (1977) Thename Danaeaceae has been found to predateMarattiaceae (Appendix 4) however Marattiaceae isbeing proposed for conservation by Murdock amp al(subm) and we maintain its usage in the usual broadsense

Terrestrial (rarely epipetric) pantropical fossils be-ginning in Carboniferous (Collinson 1996) Charactersroots large fleshy with polyarch xylem root hairs sep-tate roots stems and leaves with mucilage canals rhi-zomes fleshy short upright or creeping with a polycy-


709

clic dictyostele vernation circinate leaves large fleshy1ndash3-pinnate (rarely simple in Danaea or 3ndash5-foliate inChristensenia) with enlarged fleshy starchy stipules atthe base and swollen pulvinae along petioles and rachis-es (and sometimes other axes) petiole and stem xylempolycyclic stems and blades bearing scales pneuma-thodes (lenticels) scattered all around petioles andor ra-chises sporangia free or in round or elongate synangia(fused sporangia) lacking an annulus enclosing 1000ndash7000 spores spores usually bilateral or ellipsoid mono-lete gametophytes green surficial x = 40 (39)

IV CLASS POLYPODIOPSIDA [= FI-LICOPSIDA]

E ORDER OSMUNDALES5 Family Osmundaceae mdash Three genera Lep-

topteris Osmunda Todea Ca 20 spp monophyletic(Hasebe amp al 1995 Yatabe amp al 1999 Pryer amp al2001a 2004b) Fossils from Permian temperate andtropical Characters stem anatomy distinctive an ecto-phloic siphonostele with a ring of discrete xylemstrands these often conduplicate or twice conduplicate incross-section stipules at bases of petioles leaves dimor-phic or with fertile portions dissimilar to sterile sporan-gia large with 128ndash512 spores opening by an apical slitannulus lateral spores green subglobose trilete gameto-phytes large green cordate surficial x = 22

F ORDER HYMENOPHYLLALES6 Family Hymenophyllaceae mdash Filmy ferns

incl Trichomanaceae Nine genera (Ebihara amp al2006) two major clades (Pryer amp al 2001b) ldquotrichoma-noidrdquo and ldquohymenophylloidrdquo roughly corresponding tothe classical genera Trichomanes sl and Hymenophyl-lum sl Ca 600 spp monophyletic (Hasebe amp al 1995Dubuisson 1996 1997 Pryer amp al 2001b 2004b Ebi-hara amp al 2002 in press Dubuisson amp al 2003 Hen-nequin amp al 2003) Several segregate and monotypicgenera are nested within Hymenophyllum sl Cardio-manes Hymenoglossum Rosenstockia and Serpyllopsis(Ebihara amp al 2002 in press Hennequin amp al 2003)Several other classically defined hymenophylloid genera(subgenera) may not be monophyletic eg Mecodiumand Sphaerocionium (Hennequin amp al 2003 Ebihara ampal 2006) Microtrichomanes appears to be polyphyletic(Ebihara amp al 2004) Trichomanes sl comprises eightmonophyletic groups that are regarded here as generaAbrodictyum sl Callistopteris Cephalomanes Crepi-domanes Didymoglossum Polyphlebium Trichomanesss and Vandenboschia several of these have been sub-divided into putatively monophyletic subgenera and sec-tions (Ebihara amp al in press) Terrestrial and epiphyticpantropical and south-temperate but gametophytes sur-

vive in north-temperate regions as far north as AlaskaCharacters rhizomes slender creeping wiry or some-times erect and stouter protostelic vernation circinateblades one cell thick between veins (a few exceptions)stomata lacking cuticles lacking or highly reducedscales usually lacking on blades indument sometimes ofhairs sori marginal indusia conical (campanulate) tubu-lar or clam-shaped (bivalvate) with receptacles (at leastin trichomanoid genera) usually elongate protrudingfrom the involucres sporangia maturing graduately inbasipetal fashion each with an uninterrupted obliqueannulus spores green globose trilete gametophytes fil-amentous or ribbon-like often reproducing by fragmen-tation or production of gemmae x = 11 12 18 28 32 3334 36 and perhaps others

G ORDER GLEICHENIALES mdash IncludingDipteridales Matoniales Stromatopteridales Mo-nophyletic (Pryer amp al 2004b Schuettpelz amp al 2006)Characters root steles with 3ndash5 protoxylem poles(Schneider 1996a) antheridia with 6ndash12 narrow twistedor curved cells in walls

7 Family Gleicheniaceae mdash Gleichenioidsforking ferns incl Dicranopteridaceae Stromato-pteridaceae Six genera (Dicranopteris Diplopterygi-um Gleichenella Gleichenia Sticherus Stromatopteris)ca 125 spp monophyletic (Hasebe amp al 1995 Pryer ampal 1995 2001a 2004b) Hennipman (1996) also sugges-ted inclusion of the next two families in Gleicheniaceaehowever these are recognized here as distinct based ontheir significant morphological disparity Fossil recordbeginning in Cretaceous (Jurassic and older fossils maybelong to the Gleicheniales or represent ancestors of ex-tant Gleicheniaceae) pantropical Characters rhizomeswith a ldquovitalizedrdquo protostele or rarely solenostele leavesindeterminate blades pseudodichotomously forked (ex-cept Stromatopteris) veins free sori abaxial not mar-ginal with 5ndash15 sporangia each with a transverse-oblique annulus exindusiate round with 128ndash800spores sporangia maturing simultaneously within sorispores globose-tetrahedral or bilateral gametophytesgreen surficial with club-shaped hairs x = 22 34 3943 56

8 Family Dipteridaceae mdash Including Cheiro-pleuriaceae Two genera Cheiropleuria and Dipterisfrom India southeast Asia eastern and southern Chinacentral and southern Japan and Malesia to Melanesiaand western Polynesia (Samoa) ca 11 spp mono-phyletic (Kato amp al 2001 Pryer amp al 2004b) Fossilrecord beginning in upper Triassic Characters stemslong-creeping solenostelic or protostelic covered withbristles or articulate hairs petioles with a single vascularbundle proximally and polystelic distally blades (sterileones at least) cleft into two or often more subequal partsveins highly reticulate with included veinlets sori exin-


710

dusiate discrete compital (served by many veins) scat-tered over the surface or leaves dimorphic and the fertileones covered with sporangia sporangia maturing simul-taneously or maturation mixed with a 4-seriate stalkannuli almost vertical or slightly oblique spores ellip-soid and monolete or tetrahedral and trilete 64 or 128per sporangium gametophytes cordate-thalloid x = 33Dipteris differs from Cheiropleuria primarily in havingbilateral monolete spores (tetrahedral and trilete inCheiropleuria) and monomorphic leaves with discretesori (sporangia acrostichoid in Cheiropleuria)

9 Family Matoniaceae mdash Matonioids Twogenera (Matonia Phanerosorus) each with two sppmonophyletic sister to Dipteridaceae (Kato amp Setogu-chi 1998 Pryer amp al 2004b Schuettpelz amp al 2006)Malesia-Pacific Basin fossil record beginning in mid-Mesozoic Characters stems solenostelic with at leasttwo concentric vascular cylinders (polycyclic) and a cen-tral vascular bundle blades flabellate (Matonia) un-evenly dichotomously branched or with dichotomouspinnae veins free or slightly anastomosing around sorisori with peltate indusia sporangia maturing simultane-ously with very short stalks and oblique annuli sporesglobose-tetrahedral trilete gametophytes green thal-loid with ruffled margins antheridia large many-celledx = 26 (Matonia) 25 (Phanerosorus)

H ORDER SCHIZAEALES mdash Monophyletic(Hasebe amp al 1995 Pryer amp al 2001a 2004b Skog ampal 2002 Wikstroumlm amp al 2002) Fossil record beginningin the Jurassic (Collinson 1996) Characters fertile-ster-ile leaf blade differentiation absence of well-definedsori sporangia each with a transverse subapical contin-uous annulus

10 Family Lygodiaceae mdash Climbing ferns Asingle genus (Lygodium) ca 25 spp monophyletic(Skog amp al 2002 Wikstroumlm amp al 2002) Terrestrialpantropical Characters rhizomes creeping slender pro-tostelic bearing hairs leaves indeterminate climbingalternately pinnate primary blade divisions (pinnae)pseudodichotomously forking with a dormant bud in theaxils veins free or anastomosing sori on lobes of theultimate segments sporangia abaxial solitary one persorus each sporangium covered by an antrorse indusi-um-like subtending flange spores 128ndash256 per spo-rangium tetrahedral and trilete gametophytes greencordate surficial x = 29 30

11 Family Anemiaceae mdash Including Mohria-ceae One genus (Anemia incl Mohria) ca 100+ sppmonophyletic (Skog amp al 2002 Wikstroumlm amp al 2002)Terrestrial primarily New World but a few spp in Afri-ca India and islands in Indian Ocean Characters rhi-zomes creeping to suberect bearing hairs leaves deter-minate mostly hemidimorphic or dimorphic veins freedichotomous occasionally casually anastomosing spo-

rangia usually on a basal pair (sometimes more than twopinnae or all pinnae modified and fertile) of skele-tonized highly modified often erect pinnae spores128ndash256 per sporangium tetrahedral with strongly par-allel ridges (Tryon amp Lugardon 1991) gametophytesgreen cordate surficial x = 38

12 Family Schizaeaceae mdash Two genera (Ac-tinostachys Schizaea) ca 30 spp monophyletic (Skogamp al 2002 Wikstroumlm amp al 2002) The CretaceousSchizaeopsis is the oldest fossil assigned to this lineage(Wikstroumlm amp al 2002) Terrestrial pantropical Charac-ters blades simple (linear) or fan-shaped variously cleftand with dichotomous free veins sporangia on marginalelaminate branched or unbranched projections at bladetips not in discrete sori exindusiate spores bilateralmonolete 128ndash256 per sporangium gametophytes greenand filamentous (Schizaea) or subterranean and non-green tuberous (Actinostachys) and non-green a puz-zling array of base chromosome numbers x = 77 94103

I ORDER SALVINIALES mdash Water ferns he-terosporous ferns incl ldquoHydropteridalesrdquo Marsil-eales Pilulariales Monophyletic (Hasebe amp al 1995Pryer 1999 Pryer amp al 2001a 2004b) Characters fer-tile-sterile leaf blade differentiation veins anastomosingaerenchyma tissue often present in roots shoots andpetioles annulus absent plants heterosporous sporeswith endosporous germination monomegaspory game-tophytes reduced

13 Family Marsileaceae mdash Clover fernsincl Pilulariaceae Three genera (Marsilea PilulariaRegnellidium) ca 75 total spp monophyletic (Hasebeamp al 1995 Pryer 1999 Nagalingum amp al unpubl)Hennipman (1996) included both Salviniaceae and Azol-laceae within Marsileaceae but the spores of Marsileace-ae differ markedly from those of Salviniaceae and Azol-laceae (Schneider amp Pryer 2002) Rooted aquatics inponds shallow water or vernal pools with floating oremergent leaf blades subcosmopolitan Charactersstems usually long-creeping slender often bearing hairsleaflets 4 2 or 0 per leaf veins dichotomously branchedbut often fusing toward their tips sori borne in stalkedbean-shaped sporocarps (Nagalingum amp al in press)these arising from the rhizomes or from the base of thepetioles one to many per plant heterosporous micro-spores globose trilete megaspores globose each with anacrolamella positioned over the exine aperture(Schneider amp Pryer 2002) perine gelatinous x = 10(Pilularia) 20 (Marsilea)

14 Family Salviniaceae mdash Floating fernsmosquito ferns incl Azollaceae Two genera (Sal-vinia Azolla) ca 16 spp monophyletic (Pryer amp al1995 2004b Reid amp al 2006) Some authors separatethe genera into two families (Schneller in Kubitzki


711

1990) Plants free-floating subcosmopolitan fossil re-cord beginning in Cretaceous (Collinson 1996) Charac-ters roots present (Azolla) or lacking (Salvinia) stemsprotostelic dichotomously branched leaves sessile al-ternate small (ca 1ndash25 mm long) round to oblong en-tire veins free (Azolla) or anastomosing (Salvinia)spores of two kinds (plants heterosporous) large mega-spores and small microspores these globose triletespore germination endosporic x = 9 (Salvinia) the low-est base chromosome number known in ferns 22(Azolla)

J ORDER CYATHEALES mdash Tree ferns inclDicksoniales Hymenophyllopsidales Loxomatales Me-taxyales Plagiogyriales (Hasebe amp al 1995 Wolf amp al1999 Pryer amp al 2004b) Existing molecular evidenceindicates a close relationship among the included fami-lies The order is without obvious defining morphologi-cal characters some of the species have trunk-like stemsbut others have creeping rhizomes some have only hairson the stems and blades others have scales sori are ab-axial or marginal either indusiate or exindusiate sporesare globose or tetrahedral-globose with a trilete scar ga-metophytes green cordate

15 Family Thyrsopteridaceae mdash One genusThyrsopteris with a single species T elegans endemicto the Juan Fernaacutendez Islands clearly related to treeferns but of uncertain phylogenetic position within thisgroup (Korall amp al 2006) Characters rhizomes as-cending to erect solenostelic bearing runners clothedwith stiff pluricellular hairs leaves large 2ndash35 m longblades 3ndash5-pinnate partially dimorphic (sori oftenrestricted to proximal segments) blade axes adaxiallygrooved veins free sori terminal on the veins the outerand inner indusia fused to form asymmetric cup-likestructures each sorus with a columnar clavate recepta-cle sporangia with oblique annuli spores globose-tetra-hedral with prominent angles x = ca 78

16 Family Loxomataceae mdash Two genera (Lo-xoma Loxsomopsis) each with a single sp monophylet-ic (Pryer amp al 2001a 2004b Lehnert amp al 2001Korall amp al 2006) South American Andes southernCentral America and New Zealand Characters rhi-zomes long-creeping solenostelic bearing hairs with acircular multicellular base blades bipinnate or more di-vided veins free forked indument of uniseriate (Loxso-mopsis) to pluriseriate (Loxoma) bristles sori marginalterminal on veins each with an urceolate indusium andelongate often exserted receptacle sporangia on thickshort stalks with a slightly oblique annulus spores tetra-hedral trilete gametophytes with scalelike hairs (occur-ring also in some Cyatheaceae) x = 46 (Loxsomopsis)50 (Loxoma)

17 Family Culcitaceae mdash One genus Culcitawith two species monophyletic (Korall amp al 2006) Sis-

ter to Plagiogyriaceae and not closely related toCalochlaena with which Culcita has often been associ-ated This separation is supported by anatomical charac-ters (White amp Turner 1988 Schneider 1996a) Terrestri-al Azores Madeira Tenerife southwestern Europe andthe Neotropics Characters rhizomes creeping or ascend-ing solenostelic bearing articulate hairs petioles incross-section each with gutter-shaped vascular bundleblades large 4ndash5-pinnate-pinnatifid sparingly hairyveins free often forked sori to 3 mm wide terminal onveins paraphysate outer indusia scarcely differentiatedfrom the laminar tissue inner noticeably modifiedspores tetrahedral-globose trilete x = 66

18 Family Plagiogyriaceae mdash A single genus(Plagiogyria) with ca 15 spp (Zhang amp Nooteboom1998) monophyletic (Korall amp al 2006) Charactersstems creeping to usually erect lacking hairs or scalesleaves dimorphic blades pectinate to 1-pinnate veinssimple to 1-forked free or in fertile blades somewhatanastomosing at their ends young leaves densely cov-ered with pluricellular glandular mucilage-secretinghairs sori exindusiate sporangia borne on distal parts ofveins seemingly acrostichoid sporangial stalks 6-rowed annuli slightly oblique continuous spores tetra-hedral trilete gametophytes green cordate x = 66

19 Family Cibotiaceae Korall stat nov Basedon a full and direct reference to the Latin descriptionassociated with subfam Cibotioideae Nayar Taxon 19234 1970 ndash Type Cibotium Kaulf Jahrb Pharm 2153 1820

One genus (Cibotium) ca 11 species monophyleticwith some affinity to Dicksoniaceae as circumscribedhere (Korall amp al 2006) Terrestrial amphipacific (east-ern Asia Malesia Hawaii southern Mexico and CentralAmerica) Characters rhizomes massive creeping toascending or erect (to 6 m) solenostelic or dictyostelicbearing soft yellowish hairs at apices and persistent peti-olar bases fronds monomorphic mostly 2ndash4 m longpetioles hairy at bases with three corrugated vascularbundles arranged in an omega-shape blades large bipin-nate to bipinnate-pinnatifid or more divided secondaryand tertiary blade axes adaxially ridged veins free sim-ple or forked to pinnate stomata with 3 subsidiary cellssori marginal at vein ends indusia bivalvate each with astrongly differentiated non-green outer indusium and asimilarly modified tongue-like inner indusium paraphy-ses filiform spores globose-tetrahedral with prominentangles and a well-developed equatorial flange antheridi-al walls 5-celled x = 68

The spores of Cibotiaceae are unlike those of allother families in Cyatheales (Gastony 1982 Tryon ampLugardon 1991) with a prominent equatorial flange andwith usually thick bold plusmn parallel sometimes anasto-mosing ridges on the distal face these ridges are the


712

result of a coarsely ridged exospore which is overlain bya thin sometimes granulate perispore The spores ofLophosoria also have a prominent equatorial flange butlack distal ridges (Tryon amp Tryon 1982 Tryon amp Lu-gardon 1991) As far as is known the chromosome basenumber of x = 68 for Cibotium is also unique in the treefern clade The Hawaiian species have been extensivelystudied by Palmer (1994)

20 Family Cyatheaceae mdash Cyatheoids scalytree ferns incl Alsophilaceae HymenophyllopsidaceaeCa five genera Alsophila (incl Nephelea) Cyathea (inclCnemidaria Hemitelia Trichipteris) GymnosphaeraHymenophyllopsis Sphaeropteris (incl Fourniera)600+ spp monophyletic together with DicksoniaceaeMetaxyaceae and Cibotiaceae constituting the ldquocore treefernsrdquo (Korall amp al 2006) Several studies have ad-dressed relationships within Cyatheaceae (Conant amp al1995 1996 Hasebe amp al 1995 Stein amp al 1996 Lantzamp al 1999 Conant amp Stein 2001) and circumscriptionsof genera have varied widely (eg Tryon 1970 Holttumamp Edwards 1983) Several of these studies show con-vincingly that Cnemidaria nests within Cyathea (Conantamp al 1995 Conant amp Stein 2001 Korall amp al 2006)Hennipman (1996) included all other families hereplaced in Cyatheales (excepting Hymenophyllopsidace-ae unplaced in his ldquoconsensusrdquo classification) in his Cy-atheaceae Hymenophyllopsis (ca eight species) has thinleaves lacking stomates and is confined to the sandstonetepuis of the Guayana Shield (eastern Venezuela Guya-na northern Brazil) It has nearly always been regardedas an isolated genus in its own family (eg by Copeland1947 Tryon amp Tryon 1982) In the analysis by Wolf ampal (1999) a close relationship of Hymenophyllopsis toCyatheaceae was suggested based on a small taxonomicsampling A larger sampling by Korall (in prep) indi-cates that Hymenophyllopsis as well as Cnemidaria andTrichipteris all nest within Cyathea and together form awell supported neotropical clade The spores of Hymeno-phyllopsis are remarkably similar to some species ofCyathea (compare eg Figs 148ndash1411 with 2615ndash2618 in Tryon amp Tryon 1982) Characters associatingHymenophyllopsis with Cyatheaceae include the pres-ence of true scales on the rhizomes petiole bases andsometimes on the blades Mostly arborescent pantropi-cal fossils beginning in Jurassic or early CretaceousCharacters stems with polycyclic dictyosteles apices(and usually petiole bases) covered with large scalessometimes also with trichomidia (scurf = small scales) orhairs leaves usually large (to 5 m) petioles with obvi-ous usually discontinuous pneumathodes in two linesblades 1ndash3-pinnate (rarely simple) veins simple toforked free rarely anastomosing (mostly in some Cy-athea) sori superficial (abaxial) or terminal on the veinsand marginal or submarginal (Hymenophyllopsis) round

exindusiate or indusia saucer-like cup-like or globoseand completely surrounding sporangia or bivalvate(Hymenophyllopsis) sporangia maturing gradately withoblique annuli receptacles raised paraphyses usuallypresent spores tetrahedral trilete variously ornamentedgametophytes green cordate x = 69 (Hymenophyllopsisnot yet counted)

21 Family Dicksoniaceae nom cons mdash(Dicksonioids incl Lophosoriaceae) Three genera Cal-ochlaena Dicksonia Lophosoria) Ca 30 spp mono-phyletic (Korall amp al 2006) Terrestrial eastern AsiaAustralasia Neotropics St Helena Characters mostlyarborescent or with erect or ascending rhizomes rhi-zomes with polycyclic dictyosteles or solenostelic(Calochlaena) stem apices and usually petiolar basescovered with uniseriate hairs blades large 2ndash3 pinnateveins simple to forked free sori abaxial and exindusiate(Lophosoria) or marginal (Calochlaena Dicksonia) andeach with a bivalvate or cup-like indusium the adaxial(outer) valve formed by the reflexed segment margin andoften differently colored sporangia with oblique annulireceptacles raised paraphyses often present filiformspores globose or tetrahedral trilete x = 56(Calochlaena) 65 (Dicksonia Lophosoria)

Lophosoria (3 spp) is distinctive in having sporeswith a prominent subequatorial flange with the proximalface coarsely tuberculate the distal face perforate It hasoften been recognized as comprising its own familyLophosoriaceae (Pichi Sermolli 1977 Tryon amp Tryon1982 Kramer in Kubitzki 1990)

22 Family Metaxyaceae mdash A single genus (Me-taxya) 2 spp monophyletic (Smith amp al 2001) Terres-trial Neotropics Characters rhizomes short-creeping toascending dorsiventral solenostelic apices coveredwith pluricellular hairs petioles each with an omega-shaped corrugated vascular bundle blades simply pin-nate veins free simple or forked at the base plusmn parallelsori abaxial round scattered in several ill-defined rowsoften with several sori on the same vein with numerousfiliform paraphyses exindusiate sporangia maturingsimultaneously sporangial stalks 4-rowed annuli verti-cal or slightly oblique spores 64 per sporangium glo-bose trilete x = 95 96

K ORDER POLYPODIALES mdash Including ldquoAs-pidialesrdquo Aspleniales Athyriales Blechnales ldquoDavallia-lesrdquo Dennstaedtiales Dryopteridales Lindsaeales Lon-chitidales Monachosorales Negripteridales ParkerialesPlatyzomatales Pteridales Saccolomatales Thelypteri-dales Monophyletic (Hasebe amp al 1995 Pryer amp al1995 2001a 2004b Schneider amp al 2004c) Charac-ters indusia laterally or centrally attached (indusia lost inmany lineages) sporangial stalks 1ndash3 cells thick oftenlong sporangial maturation mixed sporangia each witha vertical annulus interrupted by the stalk and stomium


713

gametophytes green usually cordate (sometimes ribbon-shaped in some epiphytes) surficial

23 Family Lindsaeaceae mdash Lindsaeoids inclCystodiaceae Lonchitidaceae Ca eight genera Cystodi-um Lindsaea Lonchitis Odontosoria Ormoloma1 Sphe-nomeris Tapeinidium Xyropteris1 in an unpublishedthesis Barcelona (2000) advocated the establishment ofthree additional genera allied to Odontosoria and Sphe-nomeris Ca 200 spp most likely monophyletic (Wolfamp al 1994 Pryer amp al 2004b Korall amp al in pressSchuettpelz amp al in press) The inclusion of Lonchitis(traditionally associated with dennstaedtioid ferns) inLindsaeaceae is puzzling on morphological grounds butmolecular evidence strongly suggests it belongs with thelindsaeoid ferns Terrestrial or infrequently epipetric orepiphytic pantropical Characters roots with sclerenchy-matous outer cortex combined with an innermost corticallayer six cells wide (Schneider 1996a) (excepting Lon-chitis and Cystodium) rhizomes short- to long-creepingprotostelic with internal phloem or in a few taxa soleno-stelic bearing generally narrow basally attached non-clathrate scales or uniseriate hairs blades 1ndash3-pinnate ormore divided generally glabrous veins usually free for-king occasionally anastomosing without included vein-lets sori marginal or submarginal indusiate indusia open-ing towards the margin (extrorse) sometimes also at-tached at the sides or sori covered by the reflexed seg-ment margin (Lonchitis) spores tetrahedral trilete infre-quently bilateral monolete gametophytes green cor-date x = 34 38 39 44 47 48 49 50 51 perhaps oth-ers

The position of Cystodium is clearly among Poly-podiales and not Dicksoniaceae (in Cyatheales) where ithas historically been placed eg by Kramer in Kubitzki1990 Stevenson amp Loconte 1996 (Korall amp al inpress) Croft (1986) discussed its differences from dick-sonioids and recognized it at family rank A relationshipto other lindsaeoids is suggested by the molecular evi-dence and this is reflected in our classificationHowever expanded taxon sampling within early-diverg-ing lineages of Polypodiales is necessary to confirm thisor to determine whether recognition of a monotypic fam-ily Cystodiaceae is warranted (Korall amp al in press)

24 Family Saccolomataceae mdash One genus ca12 spp apparently monophyletic but more sampling isneeded to determine whether the Old World species arecongeneric with those from the New World The rela-tionships of Saccoloma (incl Orthiopteris) have beencontentious Kramer (in Kubitzki 1990) treated Saccolo-ma as a subfamily within Dennstaedtiaceae Terrestrialpantropical Characters rhizomes short-creeping to erectand somewhat trunk-like (long-creeping in most Lind-saeaceae and Dennstaedtiaceae) and dictyostelic (usuallysolenostelic in Dennstaedtiaceae protostelic with inter-

nal phloem in Lindsaeaceae) petioles each with anomega-shaped vascular strand (open end adaxial) bladespinnate to decompound lacking articulate hairs (as foundin Dennstaedtiaceae) veins free sori terminal on theveins indusia pouch- or cup-shaped spores globose-tetrahedral surface with distinctive plusmn parallel branchedridges x = ca 63

25 Family Dennstaedtiaceae mdash Dennstaedt-ioids incl Hypolepidaceae Monachosoraceae Pteridia-ceae Ca 11 genera Blotiella Coptodipteris Denn-staedtia (incl Costaricia1) Histiopteris HypolepisLeptolepia Microlepia Monachosorum Oenotrichiass1 Paesia Pteridium (bracken) Ca 170 spp mono-phyletic if lindsaeoid ferns are excluded (Pryer amp al2004b Schuettpelz amp al in press) Monachosoraceaenests within Dennstaedtiaceae (Wolf amp al 1994 Wolf1995 1997 Pryer amp al 2004b Schuettpelz amp al 2006)Terrestrial sometimes scandent pantropical Charactersrhizomes mostly long-creeping often siphonostelic orpolystelic bearing jointed hairs petioles often withepipetiolar buds usually with a gutter-shaped vascularstrand (open end adaxial) blades often large 2ndash3-pin-nate or more divided indument of hairs veins freeforked or pinnate rarely anastomosing and then withoutincluded veinlets sori marginal or submarginal linear ordiscrete indusia linear or cup-like at blade margins orreflexed over sori spores tetrahedral and trilete or reni-form and monolete gametophytes green cordate x = 2629 30 31 33 34 38 46 47 48 and probably others

26 Family Pteridaceae mdash Pteroids or pterido-ids incl Acrostichaceae Actiniopteridaceae Adiantace-ae (adiantoids maidenhairs) Anopteraceae Antrophya-ceae Ceratopteridaceae Cheilanthaceae (cheilanthoids)Cryptogrammaceae Hemionitidaceae NegripteridaceaeParkeriaceae Platyzomataceae Sinopteridaceae Taeniti-daceae (taenitidoids) Vittariaceae (vittarioids shoestringferns) Ca 50 genera 950 spp Constituent genera someof them notoriously polyphyletic or paraphyletic and inneed of redefinition (eg Cheilanthes) include Acrosti-chum Actiniopteris Adiantopsis Adiantum Aleur-itopteris Ananthacorus Anetium Anogramma Antro-phyum Argyrochosma Aspidotis Astrolepis Austro-gramme Bommeria Cassebeera Ceratopteris Cero-sora1 Cheilanthes Cheiloplecton ConiogrammeCosentinia (Nakazato amp Gastony 2001) Cryptogram-ma Doryopteris Eriosorus Haplopteris HecistopterisHemionitis Holcochlaena Jamesonia Llavea MildellaMonogramma Nephopteris1 Neurocallis NotholaenaOchropteris Onychium Paraceterach ParahemionitisPellaea (Kirkpatrick unpubl) PentagrammaPityrogramma Platyloma Platyzoma PolytaeniumPteris (incl Afropteris Anopteris) PterozoniumRadiovittaria Rheopteris Scoliosorus SyngrammaTaenitis Trachypteris and Vittaria The family thus


714

defined is monophyletic (Gastony amp Rollo 1995 1998Hasebe amp al 1995 Pryer amp al 1995 Gastony ampJohnson 2001 Schneider amp al 2004c Zhang amp al2005) Pteridaceae comprises five monophyletic groupsand if it were to be formally subdivided to reflect this ateither family or subfamily rank the following taxa couldbe recognized (1) Parkeriaceae or Parkerioideae(Acrostichum and Ceratopteris) (2) Adiantaceae orAdiantoideae (Adiantum and the ten vittarioid generaCrane amp al 1995 Hasebe amp al 1995 Hennipman1996 Crane 1997 Huiet amp Smith unpubl) (3)Cryptogrammaceae (comprising Coniogramme Crypto-gramma and Llavea Zhang amp al 2005) no subfamilyname available (4) Sinopteridaceae or Cheilan-thoideae) and (5) Pteridaceae ss or Pteridoideae con-taining Pteris (perhaps not monophyletic) its immediateallies and the taenitioid ferns (Taenitis and alliesSaacutenchez-Baracaldo 2004a b)

Terrestrial epipetric or epiphytic subcosmopolitanbut most numerous in tropics and arid regionsCharacters rhizomes long- to short-creeping ascendingsuberect or erect bearing scales (less often only hairs)blades monomorphic hemidimorphic or dimorphic in afew genera simple (mostly vittarioids) pinnate or some-times pedate sometimes decompound veins free andforking or variously anastomosing and forming a reticu-late pattern without included veinlets sori marginal orintramarginal lacking a true indusium often protectedby the reflexed segment margin or sporangia along theveins sporangia each with a vertical interrupted annu-lus receptacles not or only obscurely raised spores glo-bose or tetrahedral trilete variously ornamented mostlyx = 29 30

Platyzoma sometimes recognized as an isolatedfamily is aberrant in chromosome base number (x = 38Tindale amp Roy 2002) and in having dimorphic spores(so-called ldquoincipient heterosporyrdquo Tryon 1964) butnests with other genera of Pteridaceae subfamPteridoideae (Hasebe amp al 1995 Pryer amp al 1995)

Ceratopteris (3 spp monophyletic) nests withinPteridaceae in all molecular analyses and it appears to besister to Acrostichum (Hasebe amp al 1995 Pryer amp al1995) It has a number of strong autapomorphies thatseparate it from other Pteridaceae coarsely ridged sporeswith parallel striations spores 32 or fewer per spo-rangium sporangia with ill-defined annuli aquatic habi-tat x = 38 Consequently many taxonomists have placedit in its own family Parkeriaceae (eg Copeland 1947Pichi Sermolli 1977) Many of these autapomorphies(reduced spore number loss of annulus) are probably aconsequence of the shift to aquatic habitats

The vittarioid genera include Ananthacorus Aneti-um Antrophyum Haplopteris Hecistopteris Mono-gramma Polytaenium Radiovittaria Rheopteris Scoli-

osorus and Vittaria The presence of Rheopteris in thisclade is now clear (Huiet amp Smith unpubl) Charactersinclude linear mostly simple blades sori along veins orin linear grooves clathrate stem scales presence of ldquospi-cularrdquo cells in blade epidermis (shared with a few generaof Pteridaceae eg Adiantum) Spores in the vittarioidferns are predominantly trilete but reversals to themonolete condition have occurred in Vittaria

27 Family Aspleniaceae mdash Asplenioidsspleenworts From one to ten genera (generic delimita-tion in doubt in light of all recent molecular data egvan den Heede amp al 2003 Schneider amp al 2004b2005 Perrie amp Brownsey 2005) Regardless of the clas-sification adopted a huge majority of the species are inAsplenium even if that genus is construed in a fairlystrict sense the segregate genera Camptosorus and Loxo-scaphe clearly nest within Asplenium sl or appear relat-ed to species heretofore generally placed in Asplenium(Murakami amp al 1999 Gastony amp Johnson 2001Schneider amp al 2004b) So also are the generic segre-gates Diellia (endemic to Hawaii) Pleurosorus Phylli-tis Ceterach Thamnopteris and several others littleused in recent years (Murakami amp al 1999 Pinter amp al2002 van den Heede amp al 2003 Schneider amp al2004b) One expects that the oft-recognized but still un-sampled genera Antigramma1 Holodictyum1 Schaffner-ia1 and Sinephropteris1 also nest in Asplenium Hymen-asplenium however with a different chromosome basenumber than nearly all of the other segregates as well asdistinct root characters (Schneider 1996b Schneider ampal 2004b) appears to represent the sister clade to therest of the species in the family and this name could beadopted as a well-supported segregate genus Ca 700+spp monophyletic (Murakami amp Schaal 1994 Hasebeamp al 1995 Murakami amp al 1999 Gastony amp Johnson2001 van den Heede amp al 2003 Schneider amp al2004b 2005 Perrie amp Brownsey 2005)

Terrestrial epipetric or epiphytic subcosmopolitanbut most numerous in the tropics Characters rhizomescreeping ascending or suberect bearing clathrate scalesat apices and petiole bases (and sometimes other axes)petioles with back-to-back C-shaped vascular strandsthese fusing distally into an X-shape blades monomor-phic usually lacking acicular hairs on axes andor lami-na often with microscopic clavate hairs veins pinnate orforking usually free infrequently reticulate and thenwithout included veinlets sori elongate (linear) along theveins not usually back-to-back on the same vein usual-ly with laterally attached linear indusia sporangialstalks long 1-rowed spores reniform monolete with adecidedly winged perine x = 36 (mostly) but x = 38 39in Hymenasplenium (Murakami 1995) 38 in Boniniella

28 Family Thelypteridaceae mdash Thelypteroidsor thelypteridoids incl ldquoSphaerostephanaceaerdquo Circa


715

5ndash30 genera depending on taxonomic viewpoint com-monly accepted segregates are Cyclosorus (incl Ampel-opteris1 Amphineuron1 Chingia Christella Cyclogram-ma1 Cyclosorus ss Glaphyropteridopsis GoniopterisMeniscium Menisorus1 Mesophlebion PelazoneuronPlesioneuron Pneumatopteris Pronephrium Pseudo-cyclosorus Sphaerostephanos Stegnogramma Steirop-teris Trigonospora) Macrothelypteris PhegopterisPseudophegopteris and Thelypteris (incl AmauropeltaCoryphopteris Metathelypteris Oreopteris Parathelyp-teris and Thelypteris ss) (see Holttum 1971 Smith ampCranfill 2002) Ca 950 spp monophyletic (Hasebe ampal 1995 Smith amp Cranfill 2002 Yatabe amp al 2002)Hennipman (1996) also included Blechnaceae and theathyrioid ferns in this family a definition that wouldmake Thelypteridaceae difficult or impossible to definemorphologically

Terrestrial rarely epipetric pantropical a few tem-perate Characters rhizomes creeping ascending orerect bearing scales at apices these non-clathrate usual-ly bearing acicular hairs petioles in cross-section withtwo elongate or crescent-shaped vascular bundles facingone another these uniting distally into a gutter-shapeblades monomorphic or occasionally dimorphic usuallypinnate or pinnate-pinnatifid veins pinnate free to vari-ously and usually very regularly anastomosing with orwithout included veinlets indument of acicular hyalinehairs on blades and rhizome scales sori abaxial round tooblong rarely elongate along veins with reniform indu-sia or exindusiate sporangia with 3-rowed short to longstalks spores ellipsoid monolete perine winged to spin-ulose x = 27ndash36 Indusia have been lost independently inmany lineages within the family

29 Family Woodsiaceae mdash Athyrioids ladyferns incl Athyriaceae Cystopteridaceae Circa 15 gen-era as defined here ca 700 spp nearly 85 of them inthe two main genera Athyrium and Diplazium (inclCallipteris Monomelangium) which are both probablyparaphyletic (Wang amp al 2003) Other widely recog-nized genera include Acystopteris Cheilanthopsis Cor-nopteris Cystopteris Deparia (incl Lunathyrium Dryo-athyrium Athyriopsis and Dictyodroma Sano amp al2000b) Diplaziopsis Gymnocarpium (incl Currania)Hemidictyum Homalosorus Protowoodsia1 Pseudocys-topteris Rhachidosorus and Woodsia (incl Hymenocys-tis1 see Shmakov 2003) This family has been variouslycircumscribed and its limits are still uncertain (Hasebeamp al 1995 Sano amp al 2000a b) Wang amp al (2004) di-vided the Athyriaceae (excluding woodsioid ferns) byfar the largest component in the family into five subfam-ilies Cystopteroideae Athyrioideae Deparioideae Di-plazioideae and Rhachidosoroideae As delimited herethe Woodsiaceae may be paraphyletic to the Aspleniace-ae Blechnaceae + Onocleaceae and Thelypteridaceae

but support for this paraphylymdashor alternatively for themonophyly of the family as here definedmdashis lacking inbroad analyses (Hasebe amp al 1995 Sano amp al 2000aSchneider amp al 2004c) Because of this uncertaintycombined with the morphological grounds for the recog-nition of the Woodsiaceae as here circumscribed webelieve it is premature to adopt the alternative of erecting(or resurrecting) numerous small families to house itsconstituent genera Further sampling will likely shedadditional light on this subject and the recognition ofseveral additional families may be warranted

Mostly terrestrial subcosmopolitan Characters rhi-zomes creeping ascending or erect scales at apicesthese usually non-clathrate glabrous glandular or cili-ate petioles with two elongate or crescent-shaped vascu-lar bundles facing one another these uniting distally intoa gutter-shape blades monomorphic rarely dimorphicveins pinnate or forking free uncommonly anastomos-ing and then without included veinlets sori abaxialround J-shaped or linear with reniform to linear indusiaor exindusiate spores reniform monolete perinewinged ridged or spiny mostly x = 40 41 also 31(Hemidictyum) 33 38 39 (Woodsia) and 42 (Cysto-pteris)

30 Family Blechnaceae mdash Blechnoids inclStenochlaenaceae) Currently ca nine genera recognized(Blechnum sl Brainea Doodia Pteridoblechnum Sad-leria Salpichlaena Steenisioblechnum StenochlaenaWoodwardia) Most of the existing recognized generanest within Blechnum sl and their acceptance is depen-dent upon a revised recircumscription of Blechnum slwhich is manifestly paraphyletic in its current usage(Nakahira 2000 Cranfill 2001) Ca 200 spp mono-phyletic sister to Onocleaceae (Hasebe amp al 1995Cranfill 2001 Cranfill amp Kato 2003) Woodwardia(incl Anchistea Chieniopteris Lorinseria) appears to bean early-branching member of the Blechnaceae (Cranfillamp Kato 2003) Characters rhizomes creeping ascend-ing or erect sometimes trunk-like often bearing stolonsscaly at apex (and on blades) scales non-clathrate peti-oles with numerous round vascular bundles arranged ina ring leaves monomorphic or often dimorphic veinspinnate or forking free to variously anastomosing are-oles without included veinlets on fertile leaves formingcostular areoles bearing the sori sori in chains or linearoften parallel and adjacent to midribs indusiate with lin-ear indusia opening inwardly (toward midribs) sporan-gia with 3-rowed short to long stalks spores reniformmonolete perine winged gametophytes green cordate x= 27 28 31ndash37 (Blechnum and segregates Woodward-ia) 40 (Salpichlaena)

31 Family Onocleaceae mdash Onocleoids Fourgenera Matteuccia Onoclea Onocleopsis Pentarhizidi-um 5 spp monophyletic sister to Blechnaceae (Hasebe


716

amp al 1995 Gastony amp Ungerer 1997) Family circum-scription follows Pichi Sermolli (1977) and Gastony andUngerer (1997 their tribe Onocleeae of Dryopteridace-ae) Terrestrial largely in north-temperate regions Char-acters rhizomes long- to short-creeping to ascendingsometimes stoloniferous (Matteuccia and Onocleopsis)leaves strongly dimorphic petioles with two vascularbundles uniting distally into a gutter-shape blades pin-natifid or pinnate-pinnatifid veins free or anastomosinglacking included veinlets spores reniform brownish togreen sori enclosed (sometimes tightly) by reflexed lam-inar margins also with membranous often fugacioustrue indusia x = 37 (Onoclea) 39 40 (other genera)

32 Family Dryopteridaceae mdash Dryopteroidsor dryopteridoids incl ldquoAspidiaceaerdquo BolbitidaceaeElaphoglossaceae Hypodematiaceae PeranemataceaeCirca 40ndash45 genera 1700 spp of which 70 are in fourgenera (Ctenitis Dryopteris Elaphoglossum and Poly-stichum) Genera include Acrophorus AcrorumohraAdenoderris Arachniodes Ataxipteris1 Bolbitis (inclEgenolfia) Coveniella1 Ctenitis Cyclodium Cyrtogo-nellum Cyrtomidictyum Cyrtomium (Lu amp al 2005)Didymochlaena Dryopolystichum1 Dryopsis Dryo-pteris (incl Nothoperanema Geiger amp Ranker 2005)Elaphoglossum (incl Microstaphyla1 PeltapterisRouhan amp al 2004 Skog amp al 2004) HypodematiumLastreopsis Leucostegia Lithostegia1 LomagrammaMaxonia Megalastrum Oenotrichia pp (Tindale ampRoy 2002) Olfersia Peranema PhanerophlebiaPolystichum (incl Papuapteris Plecosorus Little ampBarrington 2003) Polybotrya Polystichopsis Revwat-tsia1 (Tindale amp Roy 2002) Rumohra Stenolepia1Stigmatopteris and Teratophyllum

Terrestrial epipetric hemiepiphytic or epiphyticpantropical also with many temperate representativesCharacters rhizomes creeping ascending or erect some-times scandent or climbing with non-clathrate scales atapices petioles with numerous round vascular bundlesarranged in a ring blades monomorphic less oftendimorphic sometimes scaly or glandular uncommonlyhairy veins pinnate or forking free to variously anas-tomosing with or without included veinlets sori usuallyround indusia round-reniform or peltate (lost in severallineages) or sori exindusiate acrostichoid in a few line-ages sporangia with 3-rowed short to long stalks sporesreniform monolete perine winged x = 41 (nearly allgenera counted) rarely 40 (presumably derived)

Dryopteridaceae as defined here is almost certainlymonophyletic if Didymochlaena Hypodematium andLeucostegia are excluded (Hasebe amp al 1995 Tsutsumiamp Kato 2006) The inclusion of these three genera mayrender this family paraphyletic but they are tentativelyincluded here pending further studies to address theirprecise phylogenetic affinities Didymochlaena with a

single species has generally been associated with othermembers of the Dryopteridaceae (as here defined) Thethree closely related species of Hypodematium on theother hand have been variously treated as composing amonogeneric family Hypodematiaceae as allied to theathyrioid ferns (eg by Kramer in Kubitzki 1990 pre-sumably based on the presence of two vascular bundlesin the petiole bases) or as close to Dryopteris (egTryon amp Lugardon 1991 using evidence from sporemorphology) Leucostegia is nearly always placed inDavalliaceae (eg by Kramer 1990) because of its sim-ilar indusia and sori terminal on the veins but it differsfrom members of Davalliaceae in the terrestrial habit themore strongly verrucate spores with rugulate perispore(Tryon amp Lugardon 1991) and x = 41 (vs x = 40 in Da-valliaceae) In a molecular phylogenetic analysis bySchneider amp al (2004c) Didymochlaena and Hypode-matium were resolved as sister to one another andtogether sister to the remainder of the Eupolypods I clade(Fig 2) but support for these relationships was lackingTsutsumi amp Kato (2006) found support for a sister rela-tionship between Hypodematium and Leucostegia andalso support for these as sister to the remaining Eu-polypods I but Didymochlaena was unsampled in theiranalysis Based on these results we therefore believe itwould be premature to segregate these genera from theDryopteridaceae

Within Dryopteridaceae as defined here nestsElaphoglossum (Hasebe amp al 1995 Sano amp al 2000a)Sometimes it is included in its own family Elaphoglos-saceae eg by Pichi Sermolli (1977) with 600ndash800spp many still undescribed Elaphoglossaceae was re-garded as comprising three genera by Pichi Sermolli(1977) but Microstaphyla and Peltapteris nest withinElaphoglossum (httpwwwnybgorgbsciresmoranelaphoglossumhtml Mickel amp Atehortuacutea 1980 Rouhan ampal 2004 Skog amp al 2004) Relationships of Elapho-glossum are often considered to be with Lomariopsis(Kramer in Kubitzki 1990) but this is refuted by two un-published topologies Elaphoglossaceae narrowly de-fined is monophyletic (Skog amp al 2001 2004) but toexclude it from Dryopteridaceae ss as delimited aboverenders the latter paraphyletic Characters of Elaphoglos-sum include simple blades (usually) and dimorphicleaves with acrostichoid sporangia

Several authors have treated most of the genera with-in our concept of Dryopteridaceae as well as Tectariace-ae Woodsiaceae and Onocleaceae as comprising amuch larger family Dryopteridaceae sl with slightlyvarying circumscriptions (eg Tryon amp Tryon 1982Kramer in Kubitzki 1990 Wagner amp Smith 1993) Withsuch a broad circumscription and unless several otherwell-circumscribed families (eg Aspleniaceae Blech-naceae Polypodiaceae Thelypteridaceae) are included


717

Dryopteridaceae is certainly paraphyletic33 Family Lomariopsidaceae mdash Lomariop-

sids incl Nephrolepidaceae sword ferns Four generaCyclopeltis Lomariopsis Nephrolepis and Thysanosori-a1 ca 70 species Characters rhizomes creeping orsometimes climbing (plants hemiepiphytic) petioleswith round vascular bundles arranged in a gutter-shapeblades 1-pinnate pinnae entire or crenate often articu-late auriculate in some genera veins free plusmn parallel orpinnate sori discrete round and with round-reniform toreniform indusia or exindusiate or sporangia acrosti-choid and the fronds dimorphic spores bilateral mono-lete variously winged or ornamented x = 41 (lowernumbers known in some Lomariopsis species)

Based on published and unpublished results it ap-pears likely that these genera form a monophyleticgroup despite the fact that such an assemblage has neverbeen proposed Lomariopsidaceae (sensu Kramer inKubitzki 1990 Moran in Davidse amp al 1995) was con-strued to comprise six genera (containing ca 800+species) Bolbitis (and segregates Edanyoa Egenolfia)Elaphoglossum Lomagramma Lomariopsis Terato-phyllum and Thysanosoria1 Based on available evi-dence we place all of the aforementioned genera exceptLomariopsis (and Thysanosoria which lacks moleculardata but appears to be closely related to Lomariopsis) inthe Dryopteridaceae (see above) Nephrolepis with ca20 spp has sometimes been included in a monogenericfamily Nephrolepidaceae (Kramer in Kubitzki 1990) Ithas been resolved and supported as sister to a large cladecomprising the Tectariaceae Oleandraceae Polypodia-ceae and Davalliaceae (Hasebe amp al 1995 Schneider ampal 2004c) however Lomariopsis was not included inthese analyses When Lomariopsis is included Nephrole-pis is resolved as sister to it and these two genera inturn are strongly supported as sister to the aforemen-tioned larger clade (Tsutsumi amp Kato 2006) and there-fore to be expunged from the Dryopteridaceae Althoughwe have here decided to tentatively include Nephrolepisin the Lomariopsidaceae the monophyly of this claderequires additional scrutiny and thus Nephrolepidaceaemay eventually require recognition

34 Family Tectariaceae mdash Tectarioids inclldquoDictyoxiphiaceaerdquo ldquoHypoderriaceaerdquo 8ndash15 generaAenigmopteris1 Arthropteris Heterogonium Hypoder-ris1 Pleocnemia Psammiosorus Psomiocarpa1 Pteri-drys Tectaria sl (incl Amphiblestra1 Camptodium1Chlamydogramme1 Cionidium Ctenitopsis Dictyo-xiphium Fadyenia Hemigramma Pleuroderris1 Pseu-dotectaria1 Quercifilix and perhaps other genera men-tioned above) and Triplophyllum (Holttum 1986) ca230 species most in Tectaria sl Generic limits espe-cially within Tectaria sl are still very much in doubtThe family appears monophyletic with the definition

given Including Tectariaceae within an expandedDryopteridaceae renders the latter polyphyletic CtenitisLastreopsis and several other genera here included inDryopteridaceae have often been considered closely re-lated to tectarioid ferns (Pichi Sermolli 1977 Holttum1986 Moran in Davidse amp al 1995) but molecular datasuggest otherwise (Hasebe amp al 1995) Terrestrialpantropical Characters rhizomes usually short-creepingto ascending dictyostelic bearing scales petioles notabscising with a ring of vascular bundles in cross-sec-tion blades simple pinnate or bipinnate sometimes de-compound indument of jointed usually short stubbyhairs on the axes veins and sometimes laminar tissueespecially on rachises and costae adaxially veins free oroften highly anastomosing sometimes with includedveinlets indusia reniform or peltate (lost in several line-ages) spores brownish reniform monolete variouslyornamented x = 40 (a few genera with x = 41 some dys-ploids with x = 39)

Arthropteris is apparently not closely related toOleandra as previously suggested (Kramer in Kubitzki1990) nor to Nephrolepis as suggested by Pichi Sermol-li (1977) Analyses that have included it show it to be sis-ter to tectarioid ferns (Hasebe amp al 1995 Tsutsumi ampKato 2006) Psammiosorus a monotypic genus endem-ic to Madagascar has in turn been placed close to Ar-thropteris (Kramer in Kubitzki 1990) or even within Ar-thropteris (Tryon amp Lugardon 1991 on the basis of thespore ornamentation) Therefore both Arthropteris andPsammiosorus are tentatively assigned to Tectariaceaealthough a Tectariaceae that includes them is more diffi-cult to define morphologically

35 Family Oleandraceae mdash Monogeneric ca40 spp sister to Davalliaceae + Polypodiaceae (Hasebeamp al 1995 Schneider amp al 2004 Tsutsumi amp Kato2006) Kramer (in Kubitzki 1990) included two generain addition to Oleandra Arthropteris (ca 12 spp) andPsammiosorus (monotypic) but with this broader cir-cumscription the family is clearly polyphyletic weinclude both of these genera in Tectariaceae Species areterrestrial epilithic or often secondary hemiepiphytesCharacters blades simple leaves articulate abscisingcleanly upon senescence from pronounced phyllopodiasori indusiate indusia round-reniform spores reniformmonolete x = 41

36 Family Davalliaceae mdash Davallioids exclGymnogrammitidaceae 4ndash5 genera Araiostegia Daval-lia (incl Humata Parasorus Scyphularia) DavallodesPachypleuria ca 65 spp Monophyletic sister to Poly-podiaceae (Hasebe amp al 1995 Ranker amp al 2004Schneider amp al 2004d Tsutsumi amp Kato 2005) butmore information needed Gymnogrammitis and Leuco-stegia are often included in Davalliaceae but the formerbelongs in Polypodiaceae (Schneider amp al 2002b)


718

while the latter is seemingly allied to Hypodematium(Dryopteridaceae Tsutsumi amp Kato 2005) Generic lim-its of Araiostegia Davallia and Pachypleuria relative toeach other are ill-defined and all of these genera appearto be paraphyletic or polyphyletic (Tsutsumi amp Kato2005) Paleotropics and subtropics Pacific Basin Char-acters plants epiphytic (most genera) or epipetric rhi-zomes long-creeping dictyostelic dorsiventral bearingscales old leaves cleanly abscising at petiole basesblades usually 1ndash4-pinnate (rarely simple) monomor-phic (rarely dimorphic) veins free forking or pinnateindument generally lacking on blades and axes butsometimes of articulate hairs sori abaxial inframarginalto well back from the margin plusmn round with cup-shapedto reniform or lunate indusia (rarely forming a submar-ginal coenosorus in Parasorus) sporangia with 3-rowedusually long stalks annuli vertical spores ellipsoidmonolete yellowish to tan perine various but usuallynot strongly winged or cristate gametophytes green cor-date x = 40

37 Family Polypodiaceae mdash Polygrams inclDrynariaceae Grammitidaceae (grammitids) Gymno-grammitidaceae Loxogrammaceae PlatyceriaceaePleurisoriopsidaceae Ca 56 genera ca 1200 spp Pan-tropical a few temperate Genera include AcrosorusAdenophorus (Ranker amp al 2003) Aglaomorpha (inclPhotinopteris Merinthosorus Pseudodrynaria Holosta-chyum Janssen amp Schneider 2005) Arthromeris Belv-isia Calymmodon Campyloneurum Ceradenia Christi-opteris (Kreier amp Schneider unpubl) ChrysogrammitisCochlidium Colysis Ctenopteris Dicranoglossum Dic-tymia Drynaria (Janssen amp Schneider 2005)Enterosora Goniophlebium sl Grammitis Lecan-opteris (Haufler amp al 2003) Lellingeria Lemmaphyl-lum Lepisorus (incl Platygyria) Leptochilus Loxo-gramme (incl Anarthropteris a monotype from NewZealand Kreier amp Schneider in press) MelpomeneMicrogramma (incl Solanopteris) MicropolypodiumMicrosorum Neocheiropteris (incl Neolepisorus)Neurodium Niphidium Pecluma Phlebodium Phym-atosorus Platycerium (Kreier amp Schneider 2006)Pleopeltis Polypodioides Polypodium ProsaptiaPyrrosia (incl Drymoglossum) Scleroglossum Sel-liguea (incl Crypsinus Polypodiopteris) Serpocaulon(Smith amp al in press) Synammia (Schneider amp al2006) Terpsichore Themelium Thylacopteris (Schnei-der amp al 2004a) and Zygophlebia1 Additional mono-typic genera include Caobangia1 DrymotaeniumGymnogrammitis Kontumia1 (Wu amp al 2005) Luisma1Pleurosoriopsis and Podosorus1

Polypodiaceae ss as often recognized (eg by Kra-mer in Kubitzki 1990) is paraphyletic because it ex-cludes the grammitids often segregated as Grammitida-ceae (Ranker amp al 2004 Schneider amp al 2004d) Ge-

neric boundaries need clarification and in particularPolypodium and Microsorum two of the largest assem-blages are known to be polyphyletic (Schneider amp al2004d) Certain previously misplaced genera are nowshown to be nested within Polypodiaceae eg Pleuro-soriopsis (Hasebe amp al 1995 Schneider amp al 2004d)and Gymnogrammitis (Schneider amp al 2002b) Polypo-diaceae contains large wholly Neotropical and whollyPaleotropical clades (Schneider amp al 2004d)

Mostly epiphytic and epipetric a few terrestrialpantropical Characters rhizomes long-creeping to short-creeping dictyostelic bearing scales petioles cleanlyabscising near their bases or not (most grammitids) leav-ing short phyllopodia blades monomorphic or dimor-phic mostly simple to pinnatifid or 1-pinnate (uncom-monly more divided) indument lacking or of hairs andor scales on the blade veins often anastomosing or retic-ulate sometimes with included veinlets or veins free(most grammitids) indument various of scales hairs orglands sori abaxial (rarely marginal) round to oblong orelliptic occasionally elongate or the sporangia acrosti-choid sometimes deeply embedded sori exindusiatesometimes covered by caducous scales when young (egLepisorus Pleopeltis) sporangia with 1ndash3-rowed usual-ly long stalks frequently with paraphyses on sporangiaor on receptacle spores hyaline to yellowish reniformand monolete (non-grammitids) or greenish and glo-bose-tetrahedral trilete (most grammitids) perine vari-ous usually thin not strongly winged or cristate mostlyx = 35 36 37 (25 and other numbers also known)

Within Polypodiaceae as defined above nest thegrammitid ferns (Ranker amp al 2004 Schneider amp al2004d) Tryon amp Tryon (1982) and Hennipman (1996)subsumed the grammitids in Polypodiaceae as we dohere Grammitids (ca 20 genera 600 species pantropi-cal) do share a large number of morphological synapo-morphies veins free (mostly) scales lacking on bladessetiform often dark red-brown hairs on leaves sporan-gial stalks 1-rowed spores green trilete gametophytesribbon-shaped Some genera of grammitids have nowbeen shown to be polyphyletic eg Ctenopteris Gram-mitis Micropolypodium and Terpsichore while othersare likely monophyletic eg Ceradenia MelpomeneProsaptia sl (Ranker amp al 2004)

ACKNOWLEDGEMENTSThe authors thank Ray Cranfill for comments on an early

draft of the manuscript We also thank Chie Tsutsumi andMasahiro Kato for sharing a pre-publication copy of their paperon Davalliaceae and related genera and two anonymous review-ers for providing helpful comments James Reveal kindly provid-ed comments and suggestions on nomenclatural aspects of our


719

work especially information summarized in appendices Our workwas supported in part by National Science Foundation grantsDEB-9616260 to ARS DEB-9615533 DEB-0089909 and DEB-0347840 to KMP DEB-9707087 to PGW and DEB-0408077 toES a postdoctoral fellowship from the Swedish Research Council(2003-2724) to PK and a German Science Foundation grantSCHN 7582-1 to HS

LITERATURE CITEDAngiosperm Phylogeny Group [APG II] 2003 An update of

the Angiosperm Phylogeny Group classification for theorders and families of flowering plants APG II Bot JLinn Soc 141 399ndash436

Barcelona J F 2000 Systematics of the Fern Genus Odonto-soria Sensu Lato (Lindsaeaceae) PhD thesis MiamiUniv Oxford Ohio

Bateman R M 1991 Paleobiological and phylogenetic im-plications of anatomically-preserved Archeocalamitesfrom the Dinantian of Oxroad Bay and Loch HumphreyBurn southern Scotland Palaeontographica B223 1ndash59

Beckert S Muhle H Pruchner D amp Knoop V 1999 Phy-logenies from mitochondrial DNA in cryptogams Pp 73in Stuessy T F amp Greimler J (eds) XVII InternationalBotanical Congress Vienna Abstracts book OrganizingCommittee Vienna

Bower F O 1926 The Ferns (Filicales) Vol 2 The Euspo-rangiate and Other Relatively Primitive Ferns CambridgeUniv Press London

Brummitt R K 1996 In defence of paraphyletic taxa Pp371ndash384 in Maesen L J G van der Burgt X M amp vanMedenbach var Rooy J M (eds) The Biodiversity ofAfrican Plants Proceedings XIVth AETFAT Congress22ndash27 Aug 1994 Kluwer Academic Publishers Nether-lands

Brummitt R K 1997 Taxonomy versus cladonomy a funda-mental controversy in biological systematics Taxon 46723ndash734

Ching R C 1940 On natural classification of the familyldquoPolypodiaceaerdquo Sunyatsenia 5 201ndash268

Ching R C 1978 The Chinese fern families and genera sys-tematic arrangement and historical origin Acta PhytotaxSin 16 (3) 1ndash19 16 (4) 16ndash37

Christensen C 1938 Filicinae Pp 522ndash550 in Verdoorn F(ed) Manual of Pteridology Martinus Nijhoff TheHague

Collinson M E 1996 ldquoWhat use are fossil fernsrdquomdash20 yearson with a review of the fossil history of extant pterido-phyte families and genera Pp 349ndash394 in Camus J MGibby M amp Johns R J (eds) Pteridology in Perspec-tive Royal Botanic Gardens Kew

Conant D S Raubeson L A Attwood D K Perera SZimmer E A Sweere J A amp Stein D B 1996 Phy-logenetic and evolutionary implications of combined anal-ysis of DNA and morphology in the Cyatheaceae Pp231ndash248 in Camus J M Gibby M amp Johns R J (eds)Pteridology in Perspective Royal Botanic Gardens Kew

Conant D S Raubeson L A Attwood D K amp Stein DB 1996 [ldquo1995rdquo] The relationships of Papuasian Cyathe-

aceae to New World tree ferns Amer Fern J 85 328ndash340Conant D S amp Stein D B 2001 Phylogenetic and geo-

graphic relationships of the tree ferns (Cyatheaceae) onMount Kinabalu Sabah Parks Nat J 4 25ndash43

Copeland E B 1947 Genera Filicum Chronica BotanicaWaltham Mass

Crabbe J A Jermy A C amp Mickel J T 1975 A new ge-neric sequence for the pteridophyte herbarium Fern Gaz11 141ndash162

Crane E H 1997 A revised circumscription of the genera ofthe fern family Vittariaceae Syst Bot 22 509ndash517

Crane E H Farrar D R amp Wendel J F 1996 [ldquo1995rdquo]Phylogeny of the Vittariaceae convergent simplificationleads to a polyphyletic Vittaria Amer Fern J 85283ndash305

Cranfill R B 2001 Phylogenetic Studies in the Polypodiales(Pteridophyta) with an Emphasis on the FamilyBlechnaceae PhD thesis Univ California Berkeley

Cranfill R B amp Kato M 2003 Phylogenetics biogeogra-phy and classification of the woodwardioid ferns(Blechnaceae) Pp 25ndash48 in Chandra S amp Srivastava M(eds) Pteridology in the New Millennium KluwerAcademic Publishers Dordrecht

Crisp M D Gilmore S R amp Weston P H 1999 Phylog-enetic relationships of two anomalous species of Pulte-naea (Fabaceae Mirbelieae) and description of a newgenus Taxon 48 701ndash714

Croft J R 1986 The stipe and rachis vasculature of the dick-sonioid fern Cystodium sorbifolium (Cystodiaceae) KewBull 41 789ndash803

Davidse G Sousa S M amp Knapp S (general eds) 1995Flora Mesoamericana Vol 1 Psilotaceae a Salviniaceae(Moran R C amp Riba R vol eds) Universidad NacionalAutoacutenoma de Meacutexico Meacutexico DF

Davies K L 1991 A brief comparative survey of aerophorestructure within the Filicopsida Bot J Linn Soc 197115ndash137

Des Marais D L Smith A R Britton D M amp Pryer KM 2003 Phylogenetic relationships and evolution of ex-tant horsetails Equisetum based on chloroplast DNA se-quence data (rbcL and trnL-F) Int J Pl Sci 164737ndash751

Dickason F G 1946 The ferns of Burma Ohio J Sci 46109ndash141

Diggs Jr G M amp Lipscomb B L 2002 What is the writerof a flora to do Evolutionary taxonomy or phylogeneticsystematics Sida 20 647ndash674

Dubuisson J-Y 1996 Evolutionary relationships within thegenus Trichomanes sensu lato (Hymenophyllaceae) basedon anatomical and morphological characters and a com-parison with rbcL nucleotide sequences preliminaryresults Pp 285ndash287 in Camus J M Gibby M amp JohnsR J (eds) Pteridology in Perspective Royal BotanicGardens Kew

Dubuisson J-Y 1997 rbcL sequences a promising tool forthe molecular systematics of the fern genus Trichomanes(Hymenophyllaceae) Molec Phylog Evol 8 128ndash138

Dubuisson J-Y Hennequin S Douzery E J P CranfillR B Smith A R amp Pryer K M 2003 rbcL phyloge-ny of the fern genus Trichomanes (Hymenophyllaceae)with special reference to neotropical taxa Int J Pl Sci164 753ndash761


720

Ebihara A Dubuisson J-Y Iwatsuki K Hennequin Samp Ito M In press A taxonomic revision of Hymenophyl-laceae Blumea 51

Ebihara A Hennequin S Iwatsuki K Bostock P DMatsumoto S Jaman R Dubuisson J-Y amp Ito M2004 Polyphyletic origin of Microtrichomanes (Prantl)Copel (Hymenophyllaceae) with a revision of thespecies Taxon 53 935ndash948

Ebihara A Iwatsuki K Kurita S amp Ito M 2002Systematic position of Hymenophyllum rolandi-principisRosenst or a monotypic genus Rosenstockia Copel(Hymenophyllaceae) endemic to New Caledonia ActaPhytotax Geobot 53 35ndash49

Gastony G J 1982 Spore morphology of the Dicksoniaceae2 The genus Cibotium Canad J Bot 60 955ndash972

Gastony G J amp Johnson W P 2001 Phylogenetic place-ments of Loxoscaphe thecifera (Aspleniaceae) andActiniopteris radiata (Pteridaceae) based on analysis ofrbcL nucleotide sequences Amer Fern J 91 197ndash213

Gastony G J amp Rollo D R 1995 [1996] Phylogeny andgeneric circumscriptions of cheilanthoid ferns(Pteridaceae Cheilanthoideae) inferred from rbcLnucleotide sequences Amer Fern J 85 341ndash360

Gastony G J amp Rollo D R 1998 Cheilanthoid ferns(Pteridaceae Cheilanthoideae) in the southwestern UnitedStates and adjacent Mexicomdasha molecular phylogeneticreassessment of generic lines Aliso 17 131ndash144

Gastony G J amp Ungerer M C 1997 Molecular systematicsand a revised taxonomy of the onocleoid ferns(Dryopteridaceae Onocleeae) Amer J Bot 84 840ndash849

Gastony G J amp Yatskievych G 2001 Morphologicalreassessment of molecular phylogenetic inferences in thexerically adapted cheilanthoid ferns (Pteridaceae Cheil-anthoideae) Pp 82ndash83 in Botany 2001 AlbuquerqueNew Mexico New Mexico

Geiger J M O amp Ranker T A 2005 Molecular phyloge-netics and historical biogeography of Hawaiian Dryopteris(Dryopteridaceae) Molec Phylog Evol 34 392ndash407

Grant V 2003 Incongruence between cladistic and taxonom-ic systems Amer J Bot 90 1263ndash1270

Greuter W McNeill J Barrie F R Burdet H M De-moulin V Filgueiras T S Nicolson D H Silva PC Skog J E Trehane P Turland N J amp Hawks-worth D L (eds) 2000 International Code of BotanicalNomenclature (Saint Louis Code) Adopted by the Six-teenth International Botanical Congress St Louis Mis-souri JulyndashAugust 1999 Koeltz Scientific Books Koumlnig-stein [Regnum Veg 138]

Guillon J M 2004 Phylogeny of horsetails (Equisetum)based on the chloroplast rps4 gene and adjacent noncod-ing sequences Syst Bot 29 251ndash259

Hasebe M Omori T Nakazawa M Sano T Kato M ampIwatsuki K 1994 rbcL gene sequences provide evidencefor the evolutionary lineages of leptosporangiate fernsProc Natl Acad Sci USA 91 5730ndash5734

Hasebe M Wolf P G Pryer K M Ueda K Ito MSano R Gastony G J Yokoyama J Manhart J RMurakami N Crane E H Haufler C H amp HaukW D 1995 [1996] Fern phylogeny based on rbcLnucleotide sequences Amer Fern J 85 134ndash181

Haufler C H Grammer W A Hennipman E Ranker TA Smith A R amp Schneider H 2003 Systematics of

the ant-fern genus Lecanopteris (Polypodiaceae) testingphylogenetic hypotheses with DNA sequences Syst Bot28 217ndash227

Hauk W D 1995 [1996] A molecular assessment of relation-ships among cryptic species of Botrychium subgenus Bo-trychium (Ophioglossaceae) Amer Fern J 85 375ndash394

Hauk W D Parks C R amp Chase M W 2003 Phyloge-netic studies of Ophioglossaceae evidence from rbcL andtrnL-F plastid DNA sequences and morphology MolecPhylog Evol 28 131ndash51

Hennequin S Ebihara A Ito M Iwatsuki K ampDubuisson J-Y 2003 Molecular systematics of the ferngenus Hymenophyllum sl (Hymenophyllaceae) based onchloroplastic coding and noncoding regions MolecPhylog Evol 27 283ndash301

Hennipman E 1996 Scientific consensus classification ofPteridophyta Pp 191ndash202 in Camus J M Gibby M ampJohns R J (eds) Pteridology in Perspective RoyalBotanic Gardens Kew

Hill C R amp Camus J M 1986 Evolutionary cladistics ofmarattialean ferns Bull Brit Mus (Nat Hist) Bot 14219ndash300

Holttum R E 1947 A revised classification of leptosporan-giate ferns J Linn Soc Bot 53 123ndash158

Holttum R E 1949 The classification of ferns Biol Rev 24267ndash296

Holttum R E 1971 Studies in the family ThelypteridaceaeIII A new system of genera in the Old World Blumea 1917ndash52

Holttum R E 1973 Posing the problems Pp 1ndash10 in JermyA C Crabbe J A amp Thomas B A (eds) ThePhylogeny and Classification of the Ferns AcademicPress London [Bot J Linn Soc 67(suppl 1)]

Holttum R E 1986 Studies in the genera allied to TectariaCav V Triplophyllum a new genus of Africa andAmerica Kew Bull 41 237ndash260

Holttum R E amp Edwards P 1983 The tree ferns of Mt Ro-raima and neighboring areas of the Guayana Highlandswith comments on the family Cyatheaceae Kew Bull 38155ndash188

Hoogland R D amp Reveal J L 2005 Index nominum famil-iarum plantarum vascularium Bot Rev 71 1ndash291

Janssen T amp Schneider H 2005 Exploring the evolution ofhumus collecting leaves in drynarioid ferns (Polypodiace-ae Polypodiidae) based on phylogenetic evidence PlSyst Evol 252 175ndash197

Judd W Campbell C S Kellogg E A Stevens P F ampDonoghue M J 2002 Plant Systematics A PhylogeneticApproach Ed 2 Sinauer Associates SunderlandMassachusetts

Kato M 1983 The classification of major groups of pterido-phytes J Fac Sci Univ Tokyo Sect 3 Bot 13 263ndash283

Kato M 1987 A phylogenetic classification of Ophioglossa-ceae Gard Bull Singapore 40 1ndash14

Kato M amp Setoguchi H 1998 An rbcL-based phylogenyand heteroblastic leaf morphology of Matoniaceae SystBot 23 391ndash400

Kato M Yatabe Y Sahashi N amp Murakami N 2001Taxonomic studies of Cheiropleuria (Dipteridaceae)Blumea 46 513ndash525

Kenrick P amp Crane P R 1997 The Origin and EarlyDiversification of Land Plants Smithsonian Institution


721

Press Washington DCKorall P Conant D S Schneider H Ueda K Nishida

H amp Pryer K M In press On the phylogenetic positionof Cystodium Itrsquos not a tree fernmdashitrsquos a polypod AmerFern J 96(2)

Korall P Pryer K M Metzgar J S Schneider H amp Co-nant D S 2006 Tree ferns monophyletic groups andtheir relationships as revealed by four protein-coding plas-tid loci Molec Phylog Evol 39 830ndash845

Kranz H D amp Huss V A R 1996 Molecular evolution ofpteridophytes and their relationship to seed plants evi-dence from complete 18S rRNA gene sequences Pl SystEvol 202 1ndash11

Kreier H-P amp Schneider H 2006 Phylogeny and biogeog-raphy of the staghorn fern genus Platycerium(Polypodiaceae Polypodiidae) Amer J Bot 93 217ndash225

Kreier H-P amp Schneider H In press Reinstatement ofLoxogramme dictyopteris for a New Zealand endemic fernknown as Anarthropteris lanceolata based on phylogenet-ic evidence Austral Syst Bot

Kubitzki K (ed) 1990 The Families and Genera of VascularPlants Vol 1 Pteridophytes and Gymnosperms (Vol edsKramer K U amp Green P S) Springer-Verlag Berlin

Lantz T C Rothwell G W amp Stockey R A 1999 Conan-tiopteris schuchmanii gen et sp nov and the role of fos-sils in resolving the phylogeny of Cyatheaceae sl J PlRes 112 361ndash381

Lehnert M Moumlnnich M Pleines T Schmidt-Lebuhn Aamp Kessler M 2001 The relictual fern genus Loxsomop-sis Amer Fern J 91 13ndash24

Lideacuten M Oxelman B Backlund A Andersson LBremer B Eriksson R Moberg R Nordal IPersson K Thulin M amp Zimmer B 1997 Charlie isour darling Taxon 46 735ndash738

Little D P amp Barrington D S 2003 Major evolutionaryevents in the origin and diversification of the fern genusPolystichum (Dryopteridaceae) Amer J Bot 90 508ndash514

Lu J-M Li D-Z Gao L-M amp Cheng X 2005 Paraphylyof Cyrtomium (Dryopteridaceae) evidence from rbcL andtrnL-F sequence data J Pl Res 118 129ndash135

Manhart J R 1994 Phylogenetic analysis of green plantrbcL sequences Molec Phylog Evol 3 114ndash127

Manhart J R 1996 [ldquo1995rdquo] Chloroplast 16S rDNAsequences and phylogenetic relationships of fern allies andferns Amer Fern J 85 182ndash192

Mehra P N 1961 Cytological evolution in ferns with partic-ular reference to Himalayan forms Proc 48th Indian SciCongr Assoc 2 1ndash24

Mickel J T 1974 Phyletic lines in modern ferns AnnMissouri Bot Gard 61 474ndash482

Mickel J T amp Atehortuacutea L G 1980 Subdivision of thegenus Elaphoglossum Amer Fern J 70 47ndash68

Moore G 1998 A comparison of traditional and phylogeneticnomenclature Taxon 47 561ndash579

Murakami N 1995 Systematics and evolutionary biology ofthe fern genus Hymenasplenium (Aspleniaceae) J PlRes 108 257ndash268

Murakami N Nogami S Watanabe M amp Iwatsuki K1999 Phylogeny of Aspleniaceae inferred from rbcLnucleotide sequences Amer Fern J 89 232ndash243

Murakami N amp Schaal B A 1994 Chloroplast DNA varia-

tion and the phylogeny of Asplenium sect Hymenaspleni-um (Aspleniaceae) in the New World tropics J Pl Res107 245ndash251

Murdock A 2005 Molecular Evolution and Phylogeny of Ma-rattioid Ferns an Ancient Lineage of Land Plants httpwww2005botanyconferenceorgenginesearchindexphp

Nagalingum N S Schneider H amp Pryer K M In pressComparative morphology of reproductive structures inheterosporous water ferns and a re-evaluation of thesporocarp Int J Plant Sci

Nakahira Y 2000 A Molecular Phylogenetic Analysis of theFamily Blechnaceae Using the Chloroplast Gene rbcLMS thesis Graduate School of Science Univ TokyoTokyo

Nakazato T amp Gastony G J 2001 Molecular phylogeneticsof Anogramma species and related genera (PteridaceaeTaenitidoideae) P 86 in Botany 2001 Albuquerque NewMexico New Mexico

Nayar B K 1970 A phylogenetic classification of the homo-sporous ferns Taxon 19 229ndash236

Olmstead R G DePamphilis C W Wolfe A D YoungN D Elisons W J amp Reeves P A 2001 Disintegrationof the Scrophulariaceae Amer J Bot 88 348ndash361

Pahnke J Goremykin V Bobrova V Troitsky AAntonov A amp Martin W 1996 Utility of rDNA internaltranscribed spacer sequences from the inverted repeat ofchloroplast DNA in pteridophyte molecular phylogeneticsPp 217ndash230 in Camus J M Gibby M amp Johns R J(eds) Pteridology in Perspective Royal Botanic GardensKew

Palmer D D 1994 The Hawaiian species of Cibotium AmerFern J 84 73ndash85

Perrie L R amp Brownsey P J 2005 Insights into the bio-geography and polyploid evolution of New ZealandAsplenium from chloroplast DNA sequence data AmerFern J 95 1ndash21

Pichi Sermolli R E G 1958 The higher taxa of thePteridophyta and their classification Pp 70ndash90 inHedberg O (ed) Systematics of today Uppsala UnivAringrsskrift 1958(6) 70ndash90

Pichi Sermolli R E G 1959 Pteridophyta Pp 421ndash493 inTurrill W B (ed) Vistas in Botany Pergamon Press LtdLondon

Pichi Sermolli R E G 1965 Adumbratio Florae Aethiopicae11 Oleandraceae Webbia 20 745ndash769

Pichi Sermolli R E G 1970 A provisional catalogue of thefamily names of living pteridophytes Webbia 25 219ndash297

Pichi Sermolli R E G 1973 Historical review of the higherclassification of the Filicopsida Pp 11ndash40 in Jermy AC Crabbe J A amp Thomas B A (eds) The Phylogenyand Classification of the Ferns Academic Press London[Bot J Linn Soc 67(suppl 1)]

Pichi Sermolli R E G 1977 Tentamen pteridophytorumgenera in taxonomicum ordinem redigendi Webbia 31313ndash512

Pichi Sermolli R E G 1981 Report of the Subcommittee forfamily names of Pteridophyta Taxon 30 163ndash168

Pichi Sermolli R E G 1982 A further contribution to thenomenclature of the families of Pteridophyta Webbia 35223ndash237



722

Pichi Sermolli R E G 1993 New studies on some familynames of Pteridophyta Webbia 47 121ndash143

Pinter I Bakker F Barrett J Cox C Gibby M Hen-derson S Morgan-Richards M Rumsey F RussellS Trewick S Schneider H amp Vogel J 2002 Phylog-enetic and biosystematic relationships in four highly dis-junct polyploid complexes in the subgenera Ceterach andPhyllitis in Asplenium (Aspleniaceae) Org Diver Evol 2299ndash311

Pryer K M 1999 Phylogeny of marsileaceous ferns and rela-tionships of the fossil Hydropteris pinnata reconsideredInt J Pl Sci 160 931ndash954

Pryer K M Schneider H amp Magalloacuten S 2004a The radi-ation of vascular plants Pp 138ndash153 in Cracraft J ampDonoghue M J (eds) Assembling the Tree of LifeOxford Univ Press New York

Pryer K M Schneider H Smith A R Cranfill R WolfP G Hunt J S amp Sipes S D 2001a Horsetails andferns are a monophyletic group and the closest living rela-tives to seed plants Nature 409 618ndash622

Pryer K M Schuettpelz E Wolf P G Schneider HSmith A R amp Cranfill R 2004b Phylogeny and evolu-tion of ferns (monilophytes) with a focus on the early lep-tosporangiate divergences Amer J Bot 91 1582ndash1598

Pryer K M Smith A R Hunt J S amp Dubuisson J-Y2001b rbcL data reveal two monophyletic groups of filmyferns (Filicopsida Hymenophyllaceae) Amer J Bot 881118ndash1130

Pryer K M Smith A R amp Skog J E 1996 [ldquo1995rdquo]Phylogenetic relationships of extant ferns based on evi-dence from morphology and rbcL sequences Amer FernJ 85 205ndash282

Ranker T A Geiger J M O Kennedy S C Smith A RHaufler C H amp Parris B S 2003 Molecular phyloge-netics and evolution of the endemic Hawaiian genusAdenophorus (Grammitidaceae) Molec Phylog Evol 26337ndash347

Ranker T A Smith A R Parris B S Geiger J M OHaufler C H Straub S C K amp Schneider H 2004Phylogeny and evolution of grammitid ferns (Gram-mitidaceae) a case of rampant morphological homoplasyTaxon 53 415ndash428

Raubeson L A amp Jansen R K 1992 Chloroplast DNA evi-dence on the ancient evolutionary split in vascular landplants Science 255 1697ndash2699

Reid J D Plunkett G M amp Peters G A 2006Phylogenetic relationships in the heterosporous fern genusAzolla (Azollaceae) based on DNA sequence data fromthree noncoding regions Int J Pl Sci 167 529ndash538

Renzaglia K S Duff R J Nickrent D L amp Garbary DJ 2000 Vegetative and reproductive innovations of earlyland plants implications for a unified phylogeny PhilosTrans Ser B 355 769ndash793

Rouhan G Dubuisson J-Y Rakotondrainibe F MotleyT J Mickel J T Labat J-N amp Moran R C 2004Molecular phylogeny of the fern genus Elaphoglossum(Elaphoglossaceae) based on chloroplast non-coding DNAsequences contributions of species from the Indian Oceanarea Molec Phylog Evol 33 745ndash763

Saacutenchez-Baracaldo P 2004a Phylogenetics and biogeogra-phy of the neotropical fern genera Jamesonia and Erio-sorus (Pteridaceae) Amer J Bot 91 274ndash284

Saacutenchez-Baracaldo P 2004b Phylogenetic relationships ofthe subfamily Taenitoideae Pteridaceae Amer Fern J 94126ndash142

Sano R Takamiya M Ito M Kurita S amp Hasebe M2000a Phylogeny of the lady fern group tribePhysematieae (Dryopteridaceae) based on chloroplastrbcL gene sequences Molec Phylog Evol 15 403ndash413

Sano R Takamiya M Kurita S Ito M amp Hasebe M2000b Diplazium subsinuatum and D tomitaroanumshould be moved to Deparia according to molecular mor-phological and cytological characters J Pl Res 113157ndash163

Schneider H 1996a Vergleichende Wurzelanatomie derFarne PhD dissertation Univ Zuumlrich Shaker Aachen

Schneider H 1996b Root anatomy of Aspleniaceae and theimplications for systematics of the fern family Fern Gaz12 160ndash168

Schneider H Janssen T Hovenkamp P Smith A RCranfill R Haufler C H amp Ranker T A 2004aPhylogenetic relationships of the enigmatic Malesian fernThylacopteris (Polypodiaceae Polypodiidae) Int J PlSci 165 1077ndash1087

Schneider H Kreier H-P Wilson R amp Smith A R 2006The Synammia enigma evidence for a temperate lineageof polygrammoid ferns (Polypodiaceae Polypodiidae) insouthern South America Syst Bot 31 31ndash41

Schneider H amp Pryer K M 2002 Structure and function ofspores in the aquatic heterosporous fern familyMarsileaceae Int J Pl Sci 163 485ndash505

Schneider H Pryer K M Cranfill R Smith A R ampWolf P G 2002a Evolution of vascular plant body plansa phylogenetic perspective Pp 330ndash364 in Cronk Q CB Bateman R M amp Hawkins J A (eds) Developmen-tal Genetics and Plant Evolution Taylor amp FrancisLondon

Schneider H Ranker T A Russell S J Cranfill R Gei-ger J M O Aguraiuja R Wood K R GrundmannM Kloberdanz K amp Vogel J C 2005 Origin of theendemic fern genus Diellia coincides with the renewal ofHawaiian terrestrial life in the Miocene Proc Roy SocLondon ser B Biol Sci 272 455ndash460

Schneider H Russell S J Cox C J Bakker F Hen-derson S Gibby M amp Vogel J C 2004b Chloroplastphylogeny of asplenioid ferns based on rbcL and trnL-Fspacer sequences (Polypodiidae Aspleniaceae) and itsimplications for the biogeography Syst Bot 29 260ndash274

Schneider H Schuettpelz E Pryer K M Cranfill RMagalloacuten S amp Lupia R 2004c Ferns diversified in theshadow of angiosperms Nature 428 553ndash557

Schneider H Smith A R Cranfill R Haufler C HRanker T A amp Hildebrand T 2002b Gymnogrammitisdareiformis is a polygrammoid fern (Polypodiaceae)mdashResolving an apparent conflict between morphologicaland molecular data Pl Syst Evol 234 121ndash136

Schneider H Smith A R Cranfill R Hildebrand T EHaufler C H amp Ranker T A 2004d Unraveling thephylogeny of polygrammoid ferns (Polypodiaceae andGrammitidaceae) exploring aspects of the diversificationof epiphytic plants Molec Phylog Evol 31 1041ndash1063

Schuettpelz E Korall P amp Pryer K M In press PlastidatpA data provide improved support for deep relationshipsamong ferns Taxon 55


723

Scotland R W amp Wortley A H 2003 How many species ofseed plants are there Taxon 52 101ndash104

Shmakov A I 2003 Review of the family Woodsiaceae(Diels) Herter of Eurasia Pp 49ndash64 in Chandra S ampSrivastava M (eds) Pteridology in the New MillenniumKluwer Academic Publishers Dordrecht

Skog J E Mickel J T Moran R C Volovsek M Zim-mer E A 2004 Molecular studies of representative spe-cies in the fern genus Elaphoglossum (Dryopteridaceae)based on cpDNA sequences rbcL trnL-F and rps4-TRNSInt J Pl Sci 165 1063ndash1075

Skog J E Mickel J T Moran R amp Zimmer E A 2001Phylogeny of the fern genus Elaphoglossum based on twochloroplast genes P 87 in Botany 2001 AlbuquerqueNew MexicoNew Mexico

Skog J E Zimmer E amp Mickel J T 2002 Additional sup-port for two subgenera of Anemia (Schizaeaceae) fromdata for the chloroplast intergenic spacer region trnL-Fand morphology Amer Fern J 92 119ndash130

Smith A R 1995 [1996] Non-molecular phylogenetic hy-potheses for ferns Amer Fern J 85 104ndash122

Smith A R amp Cranfill R B 2002 Intrafamilial relation-ships of the thelypteroid ferns (Thelypteridaceae) AmerFern J 92 131ndash149

Smith A R Kreier H-P Haufler C H Ranker T A ampSchneider H In press Serpocaulon (Polypodiaceae) anew genus segregated from old Polypodium Taxon 55

Smith A R Tuomisto H Pryer K M Hunt J S amp WolfP G 2001 Metaxya lanosa a second species in the genusand fern family Metaxyaceae Syst Bot 26 480ndash486

Stein D B Conant D S amp Valinski A E C 1996 The im-plications of chloroplast DNA restriction site variation onthe classification and phylogeny of the Cyatheaceae Pp235ndash254 in Johns R J (ed) Holttum Memorial VolumeRoyal Botanic Gardens Kew

Stevenson D W amp Loconte H 1996 Ordinal and familialrelationships of pteridophyte genera Pp 435ndash467 inCamus J M Gibby M amp Johns R J (eds) Pteridologyin Perspective Royal Botanic Gardens Kew

Sun B-Y Kim M H Kim C H amp Park C-W 2001Mankyua (Ophioglossaceae) a new fern genus fromCheju Island Korea Taxon 50 1019ndash1024

Tagawa M amp Iwatsuki K 1972 Families and genera of thepteridophytes known from Thailand Mem Fac Sci KyotoUniv Ser Biol 5 67ndash88

Thorne R F 2002 How many species of seed plants arethere Taxon 51 511ndash522

Tindale M D amp Roy S K 2002 A cytotaxonomic survey ofthe Pteridophyta of Australia Austr Syst Bot 15839ndash937

Tryon A F 1964 Platyzomamdasha Queensland fern with incip-ient heterospory Amer J Bot 51 939ndash942

Tryon A F amp Lugardon B 1991 Spores of the Pteridophy-ta Springer-Verlag New York

Tryon R 1970 The classification of the Cyatheaceae ContrGray Herb 200 1ndash53

Tryon R M amp Tryon A F 1982 Ferns and Allied Plantswith Special Reference to Tropical America Springer-Verlag Berlin

Tsutsumi C amp Kato M 2005 Molecular phylogenetic studyon Davalliaceae Fern Gaz 17 147ndash162

Tsutsumi C amp Kato M In press Evolution of epiphytes in

Davalliaceae and related ferns Bot J Linn Socvan den Heede C J Viane R L L amp Chase M W 2003

Phylogenetic analysis of Asplenium subgenus Ceterach(Pteridophyta Aspleniaceae) based on plastid and nuclearribosomal ITS DNA sequences Amer J Bot 90 481ndash493

Vangerow S Teerkorn T amp Knoop V 1999 Phylogeneticinformation in the mitochondrial nad5 gene of pterido-phytes RNA editing and intron sequences Pl Biol 1235ndash243

Wagner W H Jr 1969 The construction of a classificationPp 67ndash90 in U S National Academy of ScienceSystematic Biology U S Natl Acad Sci Publ No 1692National Academy Press Washington DC

Wagner W H Jr amp Smith A R 1993 Pteridophytes ofNorth America Pp 247ndash266 in Flora of North AmericaEditorial Committee (eds) Flora of North America Northof Mexico Vol 1 Oxford Univ Press New York

Wang M-L Chen Z-D Zhang X-C Lu S-G amp ZhaoG-F 2003 Phylogeny of the Athyriaceae evidence fromchloroplast trnL-F region sequences Acta Phytotax Sin41 416ndash426

Wang M-L Hsieh Y-T amp Zhao G-F 2004 A revised sub-division of the Athyriaceae Acta Phytotax Sin 42524ndash527

White R A amp Turner M D 1988 Calochlaena a newgenus of dicksonioid ferns Amer Fern J 78 86ndash95

Wikstroumlm N Kenrick P amp Vogel J C 2002Schizaeaceae a phylogenetic approach Rev PalaeobotPalynol 119 35ndash50

Wikstroumlm N amp Pryer K M 2005 Incongruence betweenprimary sequence data and the distribution of a mitochon-drial atp1 group II intron among ferns and horsetailsMolec Phylog Evol 36 484ndash493

Wolf P G 1995 [1996] Phylogenetic analyses of rbcL andnuclear ribosomal RNA gene sequences in Dennstaedtia-ceae Amer Fern J 85 306ndash327

Wolf P G 1996 Pteridophyte phylogenies based on analysisof DNA sequences a multiple gene approach Pp203ndash215 in Camus J M Gibby M amp Johns R J (eds)Pteridology in Perspective Royal Botanic Gardens Kew

Wolf P G 1997 Evaluation of atpB nucleotide sequences forphylogenetic studies of ferns and other pteridophytesAmer J Bot 84 1429ndash1440

Wolf P G Pryer K M Smith A R amp Hasebe M 1998Phylogenetic studies of extant pteridophytes Pp 541ndash556in Soltis D E Soltis P S amp Doyle J J (eds)Molecular Systematics of Plants II DNA SequencingKluwer Academic Publishers Boston

Wolf P G Sipes S D White M R Martines M LPryer K M Smith A R amp Ueda K 1999Phylogenetic relationships of the enigmatic fern familiesHymenophyllopsidaceae and Lophosoriaceae evidencefrom rbcL nucleotide sequences Pl Syst Evol 219263ndash270

Wolf P G Soltis P S amp Soltis D E 1994 Phylogeneticrelationships of dennstaedtioid ferns evidence from rbcLsequences Molec Phylog Evol 3 383ndash392

Wu S Phan K L amp Xiang J 2005 A new genus and twonew species of ferns from Vietnam Novon 15 245ndash249

Yatabe Y Nishida H amp Murakami N 1999 Phylogeny ofOsmundaceae inferred from rbcL nucleotide sequencesand comparison to the fossil evidence J Pl Res 112


724

397ndash404Yatabe Y Watkins J E Farrar D R amp Murakami N

2002 Genetic variation in populations of the morphologi-cally and ecologically variable fern Stegnogramma pozoisubsp mollissima (Thelypteridaceae) in Japan J Pl Res115 29ndash38

Zhang G Zhang X amp Chen Z 2005 Phylogeny of cryp-togrammoid ferns and related taxa based on rbcLsequences Nordic J Bot 23 485ndash493

Zhang X-C amp Nooteboom H P 1998 A taxonomic revi-sion of Plagiogyriaceae (Pteridophyta) Blumea 43401ndash469


725

Appendix 1 Supra-ordinal names applied to extant ferns Boldface indicates names adopted in this classification refers to Kenrick amp Crane 1997Subclass Blechnidae Doweld Subdivision Ophioglossophytina DoweldSubclass Cyatheidae Doweld Class Ophioglossopsida Thomeacute Class Cyatheopsida Doweld Subclass Osmundidae DoweldSubclass Dennstaedtiidae Doweld Class Osmundopsida DoweldSubclass Equisetidae Warming Subclass Plagiogyriideae DoweldDivision Equisetophyta D H Scott Subclass Polypodiidae Cronquist Takht amp ZimmermSubdivision Equisetophytina Reveal Division Polypodiophyta Cronquist Takht amp ZimmermClass Equisetopsida C Agardh Subdivision Polypodiophytina RevealSubclass Filicidae Hook amp Arn nom illeg Class Polypodiopsida Cronquist Takht amp ZimmermDivision Filicophyta J Mackay nom illeg Subclass Psilotidae RevealClass Filicopsida C Agardh nom illeg Division Psilotophyta HeintzeSubclass Gleichenidae Doweld Subdivision Psilotophytina Tippo ex RevealClass Gleicheniopsida Doweld Class Psilotopsida D H ScottClass Hymenophyllopsida Doweld Subclass Pterididae SchmakovSubclass Marattiidae Klinge Subkingdom Pteridobiotina UnderwDivision Marattiophyta Heintze Division Pteridophyta SchimpClass Marattiopsida Doweld Superdivision Pteridophytanae DoweldSubclass Marsileidae Doweld Subdivision Pteridophytina EnglClass Marsileopsida Trevis Class Pteridopsida RitgenClass Matoniopsida Doweld Subclass Salviniidae Cronquist Takht amp Zimmerm ex RevealSubclass Matoniidae Doweld Class Salviniopsida Kamelin amp SchmakovInfradivision Moniliformopses nom illeg Subclass Schizaeidae DoweldSubclass Ophioglossidae Klinge Class Schizaeopsida DoweldDivision Ophioglossophyta Bek


726

Appendix 3 Familial names applied to extant ferns and their taxonomic disposition Family names accepted by us arein boldface Synonyms are in italics Unpublished or otherwise illegitimate names are in quotation marks An equal sign(=) is intended to indicate that we regard the first name as a heterotypic synonym of the family name we adoptAcrostichaceae Mett ex AB Frank = Pteridaceae Dicksoniaceae MR Schomb nom cons over ThyrsopteridaceaeActiniopteridaceae Pic Serm = Pteridaceae here included in CyathealesAdiantaceae Newman nom cons over Parkeriaceae = Pteridaceae Dicranopteridaceae Ching ex Doweld = GleicheniaceaeAlsophilaceae C Presl = Cyatheaceae ldquoDictyoxiphiaceaerdquo Ching nom nud = TectariaceaeAnemiaceae Link here included in Schizaeales ldquoDidymochlaenaceaerdquo Ching nom nud = Dryopteridaceae tentativelyAngiopteridaceae Feacutee ex J Bommer = Marattiaceae Dipteridaceae Seward amp E Dale here included in GleichenialesAnopteraceae Doweld = Pteridaceae Drynariaceae Ching = PolypodiaceaeAntrophyaceae Ching = Pteridaceae Dryopteridaceae Herter nom cons over Peranemataceae here ldquoAspidiaceaerdquo Burnett nom illeg = Dryopteridaceae included in PolypodialesAspleniaceae Newman here included in Polypodiales Elaphoglossaceae Pic Serm = DryopteridaceaeAthyriaceae Alston = Woodsiaceae Equisetaceae Michx ex DC here included in EquisetalesAzollaceae Wettst = Salviniaceae ldquoFilicaceaerdquo Juss nom illegBlechnaceae Newman here included in Polypodiales Gleicheniaceae C Presl here included in GleichenialesBolbitidaceae Ching = Dryopteridaceae Grammitidaceae Newman [often misspelled Grammitaceae] =Botrychiaceae Horan = Ophioglossaceae PolypodiaceaeCeratopteridaceae Underw = Parkeriaceae = Pteridaceae Gymnogrammitidaceae Ching (incl Gymnogrammaceae spellingCheilanthaceae BK Nayar = Pteridaceae variant used by some authors) = PolypodiaceaeCheiropleuriaceae Nakai = Dipteridaceae Helminthostachyaceae Ching = OphioglossaceaeChristenseniaceae Ching = Marattiaceae Hemionitidaceae Pic Serm = PteridaceaeCibotiaceae Korall here included in Cyatheales Hymenophyllaceae Mart here included in HymenophyllalesCryptogrammaceae Pic Serm = Pteridaceae Hymenophyllopsidaceae Pic Serm = CyatheaceaeCulcitaceae Pic Serm here included in Cyatheales Hypodematiaceae Ching = Dryopteridaceae tentativelyCyatheaceae Kaulf here included in Cyatheales ldquoHypoderriaceaerdquo Ching nom nud used by various authors inclCystodiaceae JR Croft = Lindsaeaceae Dickason (1946) = TectariaceaeCystopteridaceae Schmakov Hypolepidaceae Pic Serm = DennstaedtiaceaeDanaeaceae C Agardh = Marattiaceae ldquoKaulfussiaceaerdquo Campb nom illeg = MarattiaceaeDavalliaceae MR Schomb here included in Polypodiales Lindsaeaceae C Presl here included in PolypodialesDennstaedtiaceae Lotsy here included in Polypodiales Lomariopsidaceae Alston here included in Polypodiales

Appendix 2 Ordinal and subordinal names applied to extant ferns In this list an equal sign (=) indicates that we regardthe first name as a synonym of the one we adopt refers to orders adopted by Pichi Sermolli (1977) most of thesenames are not validly published Boldface indicates orders adopted in our classification A name in quotes indicatesthat it is not validly publishedOrder ldquoAspidialesrdquo used by various authors eg Order Loxomatales Pic Serm ex Reveal = Cyatheales

Pichi Sermolli (1958 1977) nom illeg = Polypodiales Order Marattiales LinkOrder Aspleniales Pic Serm ex Reveal = Polypodiales Order Marsileales Bartl = SalvinialesOrder Athyriales Schmakov = Polypodiales Order Matoniales Pic Serm ex Reveal = GleichenialesOrder Blechnales Pic Serm ex Reveal = Polypodiales Order Metaxyales Doweld = CyathealesOrder Christenseniales Doweld = Marattiales Order Monachosorales Doweld = PolypodialesOrder Cyatheales AB Frank Order Negripteridales Pic Serm ex Reveal = PolypodialesSuborder Cyatheineae Bessey Order Ophioglossales LinkOrder ldquoDavallialesrdquo nom illeg = Polypodiales Order Osmundales LinkOrder Dennstaedtiales Doweld (name also used by Pichi Suborder Osmundineae Bessey

Sermolli 1977 where it is a nom nud) = Polypodiales Order Parkeriales AB Frank = PolypodialesOrder Dicksoniales Pic Serm ex Reveal = Cyatheales Order Pilulariales Bercht amp J Presl = SalvinialesOrder Dipteridales Doweld = Gleicheniales Order Plagiogyriales Pic Serm ex Reveal = CyathealesOrder Dryopteridales Schmakov = Polypodiales Order Platyzomatales Pic Serm ex Reveal = PolypodialesOrder Equisetales DC ex Bercht amp J Presl Order Polypodiales LinkSuborder Equisetineae Rabenh Suborder Polypodiineae BesseyOrder Filicales Dumort nom illeg Order Psilotales PrantlOrder Gleicheniales Link Suborder Psilotineae EnglSuborder Gleicheniineae Bessey Order Pteridales Doweld = PolypodialesOrder ldquoHydropteridalesrdquo (Hydropterides) used by many authors Order Saccolomatales Doweld = Polypodiales

eg Copeland (1947) Kubitzki (1990) = Salviniales Order Salviniales BartlOrder Hymenophyllales AB Frank Order Schizaeales SchimpSuborder Hymenophyllineae Bessey Suborder Schizaeineae BesseyOrder Hymenophyllopsidales Pic Serm ex Reveal = Cyatheales Order Stromatopteridales Pic Serm ex Reveal = GleichenialesOrder Lindsaeales Doweld = Polypodiales Order Thelypteridales Doweld = PolypodialesOrder Lonchitidales Doweld = Polypodiales


727

Appendix 4 Familial ordinal and supra-ordinal names for extant ferns with citations from Reveal amp Doweld (subm)as revised from Hoogland amp Reveal (2005) to comply with the Vienna Code Orthographic variations in spelling in orig-inal references are in quotation marks Italicized names are not validly published refers to ordinal names used byPichi Sermolli (1977) some of them unpublished some published by earlier authorities Commonly used ldquodescriptivenamesrdquo (Art 161) eg Hydropterides are also given but no attempt has been made to include all such names or todetermine first usage no good index is available for thesebullFam Acrostichaceae Mett ex AB Frank Syn Pflanzenk ed 2 3 1453 1458 1874 nom rej in favor of AdiantaceaebullFam Actiniopteridaceae Pic Serm Webbia 17 5 31 Aug 1962 bullFam Adiantaceae Newman Hist Brit Ferns 5 1ndash5 Feb 1840 nom cons over Parkeriaceae (1825)bullFam Alsophilaceae C Presl Gefaumlssbuumlndel Farrn 22 (footnote) 32 1847bullFam Anemiaceae Link Fil Spec 23 3ndash10 Sep 1841bullFam Angiopteridaceae Feacutee ex J Bommer Bull Soc Bot Belgique 5 345 359 before 28 May 1867 (ldquoAngiopterideaerdquo)bullFam Anopteraceae Doweld Tent Syst Pl Vasc li 23 Dec 2001bullFam Antrophyaceae Ching Acta Phytotax Sin 16(3) 11 Aug 1978 ndash basionym Antrophyoideae Link Fil Spec 140 3ndash10 Sep 1841

(ldquoAntrophyaceaerdquo)bullFam Aspidiaceae Burnett Outl Bot 324 1156 Feb 1835 nom illeg (Arts 183) The name is illegitimate because it was based on an

illegitimate generic name (Art 183)Order Aspidiales nom illeg used by various authors eg Pichi Sermolli (1958 1977) does not satisfy Art 161(a) that a name be taken

from a legitimate name of an included familybullFam Aspleniaceae Newman Hist Brit Ferns 6 1ndash5 Feb 1840Order Aspleniales Pic Serm ex Reveal Phytologia 79 72 29 Apr 1996bullFam Athyriaceae Alston Taxon 5 25 25 Mar 1956Order Athyriales Schmakov Turczaninowia 4 55 15 May 2001bullFam Azollaceae Wettst Handb Syst Bot 2 77 Nov 1903bullFam Blechnaceae Newman Hist Brit Ferns ed 2 8 1844 Order Blechnales Pic Serm ex Reveal Phytologia 74 175 25 Mar 1993Subclass Blechnidae Doweld New Syllabus 368 May 2006bullFam Bolbitidaceae Ching Acta Phytotax Sin 16(3) 15 Aug 1978 ndash basionym Bolbitideae Pic Serm Webbia 23 381 30 Jun 1969bullFam Botrychiaceae Horan Char Ess Fam 15 30 Jun 1847bullFam Ceratopteridaceae Underw Our Native Ferns ed 6 65 78 JulndashAug 1900bullFam Cheilanthaceae BK Nayar Taxon 19 233 29 Apr 1970bullFam Cheiropleuriaceae Nakai Bot Mag (Tokyo) 42 210 Apr 1928bullFam Christenseniaceae Ching Bull Fan Mem Inst Biol Bot 10 227 25 Dec 1940 Order Christenseniales Doweld Tent Syst Pl Vasc vii 23 Dec 2001bullFam Cibotiaceae Korall in A R Smith amp al Taxon 55 712 2006 ndash basionym Cibotioideae B K Nayar Taxon 19 234 1970bullFam Cryptogrammaceae Pic Serm Webbia 17 299 20 Apr 1963bullFam Culcitaceae Pic Serm Webbia 24 702 28 Apr 1970bullFam Cyatheaceae Kaulf Wesen Farrenkr [119] JulndashSep 1827Order Cyatheales AB Frank Syn Pflanzenk ed 2 3 1452 1456 1877 (ldquoCyatheaceaerdquo)Subclass Cyatheidae Doweld Tent Syst Pl Vasc xii 23 Dec 2001Suborder Cyatheineae Bessey Bot High Schools 376 14ndash21 Aug 1880 (ldquoCyatheaceaerdquo)Class Cyatheopsida Doweld Tent Syst Pl Vasc xii 23 Dec 2001bullFam Cystodiaceae JR Croft Kew Bull 41 797 20 Oct 1986

Appendix 3 ContinuedLonchitidaceae Doweld = Lindsaeaceae Platyceriaceae Ching = PolypodiaceaeLophosoriaceae Pic Serm here included in Dicksoniaceae Platyzomataceae Nakai = PteridaceaeLoxogrammaceae Ching ex Pic Serm = Polypodiaceae Pleurosoriopsidaceae Kurita amp Ikebe ex Ching = PolypodiaceaeLoxomataceae C Presl [often misspelled ldquoLoxsomaceaerdquo] here Polypodiaceae J Presl here included in Polypodiales

included in Cyatheales Psilotaceae JW Griff amp Henfr here included in PsilotalesLygodiaceae M Roem here included in Schizaeales Pteridaceae EDM Kirchn here included in PolypodialesMarattiaceae Kaulf nom cons prop here included in Marat- Pteridiaceae Ching = Dennstaedtiaceae

tiales antedated by Danaeaceae (Murdock amp al subm) Saccolomataceae Doweld here included in PolypodialesMarsileaceae Mirb here included in Salviniales Salviniaceae Martynov here included in SalvinialesMatoniaceae C Presl here included in Gleicheniales Schizaeaceae Kaulf here included in SchizaealesMetaxyaceae Pic Serm here included in Cyatheales Sinopteridaceae Koidz nom rej in favor of Adiantaceae =Mohriaceae CF Reed = Anemiaceae PteridaceaeMonachosoraceae Ching = Dennstaedtiaceae ldquoSphaerostephanaceaerdquo Ching nom nud = ThelypteridaceaeNegripteridaceae Pic Serm = Pteridaceae Stenochlaenaceae Ching = BlechnaceaeNephrolepidaceae Pic Serm = Lomariopsidaceae tentatively Stromatopteridaceae Bierh = GleicheniaceaeOleandraceae Ching ex Pic Serm here included in Polypodiales Taenitidaceae Pic Serm = PteridaceaeOnocleaceae Pic Serm here included in Polypodiales Tectariaceae Panigrahi here included in PolypodialesOphioglossaceae Martynov here included in Ophioglossales Thelypteridaceae Pic Serm here included in PolypodialesOsmundaceae Martynov here included in Osmundales Thyrsopteridaceae C Presl here included in CyathealesParkeriaceae Hook = Pteridaceae Tmesipteridaceae Nakai = PsilotaceaePeranemataceae (C Presl) Ching = Dryopteridaceae Trichomanaceae Burmeist = HymenophyllaceaePilulariaceae Mirb ex DC (Pilulariae) = Marsileaceae Vittariaceae Ching = PteridaceaePlagiogyriaceae Bower here included in Cyatheales Woodsiaceae Herter here included in Polypodiales


728

Appendix 4 ContinuedbullFam Cystopteridaceae Schmakov Turczaninowia 4 60 15 May 2001 ndash basionym Cystopteridinae Payer Bot Crypt 198 1850

(ldquoCystopterideaerdquo)bullFam Danaeaceae C Agardh Aphor Bot 117 19 Jun 1822 nom rej prop in favor of Marattiaceae nom cons prop (Murdock amp al subm)bullFam Davalliaceae MR Schomb Reis Br-Guiana 2 883 12ndash15 Jul 1848 ndash basionym Davallieae Gaudich in Freyc Voy Uraniae Bot 370

18 Jul 1829Order Davalliales nom illeg used by Pichi Sermolli (1958)bullFam Dennstaedtiaceae Lotsy Vortr Bot Stammesgesch 2 655 9 Aug 1909 (ldquoDennstaedtineaceaerdquo)Order Dennstaedtiales Doweld Tent Syst Pl Vasc x 23 Dec 2001Subclass Dennstaedtiidae Doweld Tent Syst Pl Vasc x 23 Dec 2001bullFam Dicksoniaceae MR Schomb Reis Br-Guiana 2 1047 12ndash15 Jul 1848 nom cons over Thyrsopteridaceae (1847) ndash basionym

Dicksonieae Gaudich in Freycinet Voy Uranie Bot 367 18 Jul 1829Order Dicksoniales Pic Serm ex Reveal Phytologia 74 175 25 Mar 1993bullFam Dicranopteridaceae Ching ex Doweld Tent Syst Pl Vasc x 23 Dec 2001 nom invalid in Ching Acta Phytotax Sin 3 94 1954bullFam Dictyoxiphiaceae Ching Sunyatsenia 5 218 1940 nom nud without Latin descriptionbullFam Didymochlaenaceae Ching Sunyatsenia 5 253 1940 nom nud without Latin descriptionbullFam Dipteridaceae Seward amp E Dale Philos Trans ser B 194 487 499 502 Dec 1901 (ldquoDipteridinaerdquo) ndash basionym Dipteridinae Diels in

Engler amp Prantl Nat Pflanzenfam I 4 167 21 Jul 1899Order Dipteridales Doweld Tent Syst Pl Vasc x 23 Dec 2001bullFam Drynariaceae Ching Acta Phytotax Sin 16(4) 19 Nov 1978bullFam Dryopteridaceae Herter Rev Sudamer Bot 9 15 1949 nom cons over Peranemataceae ChingOrder Dryopteridales Schmakov Turczaninowia 4 66 67 15 May 2001bullFam Elaphoglossaceae Pic Serm Webbia 23 209 20 Nov 1968bullFam Equisetaceae Michx ex DC Essai Propr Meacuted Pl 49 Jul 1804Order Equisetales DC ex Bercht amp J Presl Prir Rostlin 271 JanndashApr 1820 (ldquoEquisetaceaerdquo)Subclass Equisetidae Warming Osnov Bot 221 22ndash28 Apr 1883 (ldquoEquisetinaerdquo)Suborder Equisetineae Rabenh Deutschl Krypt-Fl 2 xvi 332 JanndashFeb 1849 (ldquoEquisetaceaerdquo)Division Equisetophyta DH Scott Stud Foss Pl 13 489 492 493 Sep 1900 (ldquoEquisetalesrdquo)Subdivision Equisetophytina Reveal Phytologia 79 70 29 Apr 1996Class Equisetopsida C Agardh Classes Pl 7 20 May 1825 (ldquoEquisetaceaerdquo)bullFam Filicaceae Juss Gen Pl 14 4 Aug 1789 (ldquoFilicesrdquo) nom illeg (Art 183)Order Filicales Dumort Anal Fam Pl 67 1829 (ldquoFilicarieaerdquo) nom illeg (Art 161)Subclass Filicidae Hook amp Arn Brit Fl ed 6 1 564 Jul 1850 (ldquoFilicesrdquo) nom illeg (Art 161)Division Filicophyta J Mackay Fl Hiber 336 Jul 1836 nom illeg (Art 161)Class Filicopsida C Agardh Aphor Bot 114 19 Jul 1822 (ldquoFilicesrdquo) nom illeg (Art 161) Used by many authors including KampC 1997 for

what we here call PolypodiopsidabullFam Gleicheniaceae C Presl Reliq Haenk 1 70 JunndashNov 1825 (ldquoGleicheniaerdquo) ndash basionym [unranked] Gleicheneae R Br Prodr 160 27

Mar 1810Order Gleicheniales Link Handbuch 3 10 1833 (ldquoGleicheneaerdquo)Subclass Gleichenidae Doweld New Syllabus 337 May 2006Suborder Gleicheniineae Bessey Bot High Schools 376 14ndash21 Aug 1880 (ldquoGleicheniaceaerdquo)Class Gleicheniopsida Doweld Tent Syst Pl Vasc ix 23 Dec 2001bullFam Grammitidaceae Newman Hist Brit Ferns 7 1ndash5 Feb 1840 (ldquoGrammitideaerdquo) Often misspelled Grammitaceae see eg Pichi Sermolli

1993)bullFam Gymnogrammitidaceae Ching Acta Phytotax Sin 11 12 Jan 1966 Often misspelled GymnogrammaceaebullFam Helminthostachyaceae Ching Bull Fan Mem Inst Biol Bot 10 235 15 Mar 1941bullFam Hemionitidaceae Pic Serm Webbia 21 487 15 Jul 1966Order Hydropterides Campbell Mosses and Ferns 304 307 1895 Descriptive name (Art 161)bullFam Hymenophyllaceae Mart Consp Regni Veg 3 SepndashOct 1835 Order Hymenophyllales AB Frank Syn Pflanzenk ed 2 3 1452 1474 1877 (ldquoHymenophyllaceaerdquo)Suborder Hymenophyllineae Bessey Bot High Schools 376 14ndash21 Aug 1880 (ldquoHymenophyllaceaerdquo)Class Hymenophyllopsida Doweld Tent Syst Pl Vasc ix 23 Dec 2001bullFam Hymenophyllopsidaceae Pic Serm Webbia 24 712 28 Apr 1970Order Hymenophyllopsidales Pic Serm ex Reveal Phytologia 74 175 25 Mar 1993bullFam Hypodematiaceae Ching Acta Phytotax Sin 13(1) 96 Jan 1975bullFam Hypoderriaceae Ching Sunyatsenia 5 245 1940 nom nud without Latin descriptionbullFam Hypolepidaceae Pic Serm Webbia 24 705 28 Apr 1970bullFam Kaulfussiaceae Campb Evol Land Pl 333 1940 nom illeg (Art 183) because it is based on the illegitimate generic name Kaulfussia

Blume (1828) non Dennstedt (1818) nec Nees (1820)ndash basionym Kaulfussioideae Campb Struct Moss Ferns ed 2 298 300 Sep 1905(ldquoKaulfussieaerdquo) nom illeg See also Pichi Sermolli Webbia 12 353 1957

bullFam Lindsaeaceae C Presl in M R Schomb Reis Br-Guiana 2 883 12ndash15 Jul 1848Order Lindsaeales Doweld New Syllabus 353 May 2006bullFam Lomariopsidaceae Alston Taxon 5 25 25 Mar 1956bullFam Lonchitidaceae Doweld New Syllabus 353 May 2006Order Lonchitidales Doweld New Syllabus 353 May 2006bullFam Lophosoriaceae Pic Serm Webbia 24 700 28 Apr 1970bullFam Loxogrammaceae Ching ex Pic Serm Webbia 29 11 10 Feb 1975bullFam Loxomataceae C Presl Gefaumlssbuumlndel Farrn 31 1847 (ldquoLoxsomaceaerdquo)Order Loxomatales Pic Serm ex Reveal Phytologia 74 175 25 Mar 1993 (ldquoLoxsomalesrdquo)bullFam Lygodiaceae M Roem Handb Allg Bot 3 520 1840 (ldquoLygodieaerdquo)bullFam Marattiaceae Kaulf Enum Filic 31 8 Aprndash29 May 1824 nom cons prop over Danaeaceae (1822) (Murdock amp al subm)


729

Appendix 4 ContinuedOrder Marattiales Link Hort Berol 2 148 JulndashDec 1833 (ldquoMarattiaceaerdquo)Subclass Marattiidae Klinge Fl Est-Liv-Churland 1 93 22ndash28 Jun 1882 (ldquoMarattiaceaerdquo)Division Marattiophyta Heintze Cormofyt Fylog 22 1927Class Marattiopsida Doweld Tent Syst Pl Vasc vii 23 Dec 2001bullFam Marsileaceae Mirb in Lam amp Mirb Hist Nat Veacuteg 5 126 21 Nov 1802 (ldquoMarsileaerdquo)Order Marsileales Bartl in Martius Consp Regn Veg 4 SepndashOct 1835 (ldquoMarsileaceaerdquo)Subclass Marsileidae Doweld Tent Syst Pl Vasc x 23 Dec 2001Class Marsileopsida Trevis Bull Soc Roy Bot Belgique 16 5 after 6 May 1877 (ldquoMarsiglialesrdquo)bullFam Matoniaceae C Presl Gefaumlssbuumlndel Farrn 32 1847Order Matoniales Pic Serm ex Reveal Phytologia 74 175 25 Mar 1993Subclass Matoniidae Doweld New Syllabus 338 May 2006Class Matoniopsida Doweld Tent Syst Pl Vasc vii 23 Dec 2001bullFam Metaxyaceae Pic Serm Webbia 24 701 28 Apr 1970Order Metaxyales Doweld Tent Syst Pl Vasc xii 23 Dec 2001bullFam Mohriaceae CF Reed Bol Soc Brot ser 2 21 168 11 May 1948bullFam Monachosoraceae Ching Acta Phytotax Sin 16(4) 17 Nov 1978Order Monachosorales Doweld New Syllabus 356 May 2006Infradivision Moniliformopses Kenrick amp P Crane nom nud in Kenrick amp Crane 1997bullFam Negripteridaceae Pic Serm Nuovo Giorn Bot Ital ser 2 53 160 25 Nov 1946Order Negripteridales Pic Serm ex Reveal Phytologia 74 176 25 Mar 1993bullFam Nephrolepidaceae Pic Serm Webbia 29 8 10 Feb 1975bullFam Oleandraceae Ching ex Pic Serm Webbia 20 745 20 Dec 1965bullFam Onocleaceae Pic Serm Webbia 24 708 28 Apr 1970bullFam Ophioglossaceae Martynov Tekhno-Bot Slovar 438 3 Aug 1820 (ldquoOphioglosseaerdquo) ndash basionym [unranked] Ophioglosseae R Br Prodr

163 27 Mar 1810Order Ophioglossales Link Hort Berol 2 151 JulndashDec 1833 (ldquoOphioglossaceaerdquo)Subclass Ophioglossidae Klinge Fl Est-Liv-Churland 1 94 22ndash28 Jun 1882 (ldquoOphioglosseaerdquo)Division Ophioglossophyta Bek Kurs Bot 1 89 1863 (ldquoOphioglosseaerdquo)Subdivision Ophioglossophytina Doweld Tent Syst Pl Vasc ix 23 Dec 2001Class Ophioglossopsida Thomeacute Lehrb Bot ed 4 205 31 Mar 1874 (ldquoOphioglosseaerdquo)bullFam Osmundaceae Martynov Tekhno-Bot Slovar 445 3 Aug 1820Order Osmundales Link Hort Berol 2 138 JulndashDec 1833 (ldquoOsmundaceaerdquo)Subclass Osmundidae Doweld New Syllabus 334 May 2006Suborder Osmundineae Bessey Bot High Schools 377 14ndash21 Aug 1880 (ldquoOsmundaceaerdquo)Class Osmundopsida Doweld Tent Syst Pl Vasc ix 23 Dec 2001bullFam Parkeriaceae Hook Exot Fl 2 ad t 147 Mar 1825 nom rej in favor of AdiantaceaeOrder Parkeriales AB Frank Syn Pflanzenk ed 2 3 1452 1458 1877 (ldquoParkeriaceaerdquo)bullFam Peranemataceae Ching Sunyatsenia 5 208 246 30 Oct 1940 (ldquoPerenemaceaerdquo) nom rej in favor of Dryopteridaceae Herter (1949) ndash

basionym Peranemateae C Presl [Tent pterid] Abh Koumlnigl Boumlhm Ges Wiss ser 4 5 64 before 2 Dec 1836bullFam Pilulariaceae Mirb ex DC Essai Propr Meacuted Pl 48 Jul 1804 (ldquoPilulariaerdquo)Order Pilulariales Bercht amp J Presl Prir Rostlin 272 JanndashApr 1820 (ldquoPilulariaerdquo)bullFam Plagiogyriaceae Bower Ann Bot (London) 40 484 Apr 1926Order Plagiogyriales Pic Serm ex Reveal Phytologia 74 176 25 Mar 1993Subclass Plagiogyriidae Doweld Tent Syst Pl Vasc xii 23 Dec 2001bullFam Platyceriaceae Ching Acta Phytotax Sin 16(3) 18 Aug 1978bullFam Platyzomataceae Nakai Bull Natl Sci Mus Tokyo 29 4 Dec 1950Order Platyzomatales Pic Serm ex Reveal Phytologia 74 176 25 Mar 1993bullFam Pleurosoriopsidaceae Kurita amp Ikebe ex Ching Acta Phytotax Sin 16(4) 17 Nov 1978bullFam Polypodiaceae J Presl amp C Presl Delic Prag 159 Jul 1822Order Polypodiales Link Hort Berol 2 5 JulndashDec 1833 (ldquoPolypodiaceaerdquo)Subclass Polypodiidae Cronquist Takht amp Zimmerm Taxon 15 133 Apr 1966Suborder Polypodiineae Bessey Bot High Schools 377 14ndash21 Aug 1880 (ldquoPolypodiaceaerdquo)Division Polypodiophyta Cronquist Takht amp Zimmerm Taxon 15 133 Apr 1966Subdivision Polypodiophytina Reveal Phytologia 79 70 29 Apr 1996Class Polypodiopsida Cronquist Takht amp Zimmerm Taxon 15 133 Apr 1966bullFam Psilotaceae JW Griff amp Henfr Microgr Dict 540 1 Nov 1855 (ldquoPsiloteaerdquo)Order Psilotales Prantl Lehrb Bot ed 5 183 5 Jan 1884 (ldquoPsilotaceaerdquo)Subclass Psilotidae Reveal Phytologia 79 70 29 Apr 1996Suborder Psilotineae Engl Syllabus ed 2 64 May 1898Division Psilotophyta Heintze Cormofyt Fylog 22 1927Subdivision Psilotophytina Tippo ex Reveal Phytologia 79 70 29 Apr 1996Class Psilotopsida DH Scott Stud Foss Pl ed 2 616 631 632 May 1909bullFam Pteridaceae EDM Kirchn Schul-Bot 109 13ndash20 Oct 1831 (ldquoPteroideaerdquo) Often cited as Pteridaceae Reichb Handb Nat

Pflanzensyst 138 1837Order Pteridales Doweld Tent Syst Pl Vasc xi 23 Dec 2001bullFam Pteridiaceae Ching Acta Phytotax Sin 13(1) 96 Jan 1975Subclass Pterididae Schmakov Turczaninowia 4 42 15 May 2001Subkingdom Pteridobiotina Underw in Britton amp A Br Illus Fl N US 1 1 15 Aug 1896 (ldquoPteridophytardquo)Division Pteridophyta Schimp in Zittel Handb Palaeontol 2(1) 1 15 Sep 1879Superdivision Pteridophytanae Doweld Tent Syst Pl Vasc v 23 Dec 2001


730

Appendix 5 Index to genera with family assignments proposed in this classification All accepted genera (but not allsynonyms) in Kramer in Kubitzki (1990) are accounted for here Genera newly described or recircumscribed since 1990are also included Accepted names in roman synonyms in italics Family numbers assigned in textAbacopteris = Cyclosorus Archangiopteris = Angiopteris Ceterach = Asplenium Cyathea (20)Abrodictyum (6) Argyrochosma (26) Ceterachopsis = Asplenium Cyclodium (32)Acrophorus (32) Arthromeris (37) Cheilanthes (26) Cyclogramma = CyclosorusAcrorumohra (32) Arthropteris (34) Cheilanthopsis (29) Cyclopeltis (33)Acrosorus (37) Aspidotis (26) Cheiloplecton (26) Cyclosorus (28)Acrostichum (26) Aspleniopsis = Austrogramme Cheiroglossa = Ophioglossum Cyrtogonellum (32)Actiniopteris (26) Asplenium (27) Cheiropleuria (8) Cyrtomidictyum (32)Actinostachys (12) Astrolepis (26) Chieniopteris = Woodwardia Cyrtomium (32)Acystopteris (29) Ataxipteris (32) Chingia = Cyclosorus Cystodium (23)Adenoderris (32) Athyriopsis = Deparia Chlamydogramme = Tectaria Cystopteris (29)Adenophorus (37) Athyrium (29) Christella = Cyclosorus Danaea (4)Adiantopsis (26) Austrogramme (26) Christensenia (4) Davallia (36)Adiantum (26) Azolla (14) Christiopteris (37) Davalliopsis = TrichomanesAenigmopteris (34) Belvisia (37) Chrysochosma = Notholaena Davallodes (36)Afropteris = Pteris Blechnum (30) Chrysogrammitis (37) Dennstaedtia (25)Aglaomorpha (37) Blotiella (25) Cibotium (19) Deparia (29)Aleuritopteris (26) Bolbitis (32) Cionidium = Tectaria Diacalpe = Peranema Allantodia = Diplazium Bommeria (26) Cnemidaria = Cyathea Dicksonia (21)Alsophila (20) Botrychium (1) Cochlidium (37) Dicranoglossum (37)Amauropelta = Thelypteris Botrypus = Botrychium Colysis (37) Dicranopteris (7)Ampelopteris = Cyclosorus Brainea (30) Coniogramme (26) Dictymia (37)Amphiblestra = Tectaria Callipteris = Diplazium Coptodipteris (25) Dictyocline = CyclosorusAmphineuron = Cyclosorus Callistopteris (6) Cornopteris (29) Dictyodroma = DepariaAnanthacorus (26) Calochlaena (21) Coryphopteris = Thelypteris Dictyoxiphium = TectariaAnarthropteris = Loxogramme Calymmodon (37) Cosentinia (26) Didymochlaena (32)Anchistea = Woodwardia Camptodium = Tectaria Costaricia = Dennstaedtia Didymoglossum (6)Anemia (11) Camptosorus = Asplenium Coveniella (32) Diellia = AspleniumAnetium (26) Campyloneurum (37) Crepidomanes (6) Diplaziopsis (29)Angiopteris (4) Caobangia (37) Crypsinus = Selliguea Diplazium (29)Anogramma (26) Cardiomanes = Hymenophyllum Cryptogramma (26) Diplopterygium (7)Anopteris = Pteris Cassebeera (26) Ctenitis (32) Dipteris (8)Antigramma = Asplenium Cephalomanes (6) Ctenitopsis = Tectaria Doodia (30)Antrophyum (26) Ceradenia (37) Ctenopteris (37) Doryopteris (26)Arachniodes (32) Ceratopteris (26) Culcita (17) Drymoglossum = PyrrosiaAraiostegia (36) Cerosora (26) Currania = Gymnocarpium Drymotaenium (37)

Appendix 4 ContinuedSubdivision Pteridophytina Engl Fuumlhrer Garten Breslau 10 11 May 1886 (ldquoPteridophytardquo) Class Pteridopsida RitgenAufeinanderfolge Org Gest 63 1828 (ldquoFilices s Pteridesrdquo)bullFam Saccolomataceae Doweld New Syllabus 354 May 2006Order Saccolomatales Doweld New Syllabus 354 May 2006bullFam Salviniaceae Martynov Tekhno-Bot Slovar 559 3 Aug 1820 (ldquoSalviniaerdquo)Order Salviniales Link Hort Berol 3 155 JulndashDec 1833 (ldquoSalviniaceaerdquo)Subclass Salviniidae Pic Serm ex Reveal Phytologia 79 70 29 Apr 1996Class Salviniopsida Kamelin amp Schmakov in Kamelin Fl Altaia 253 15 May 2005bullFam Schizaeaceae Kaulf Wesen Farrenkr [119] JulndashSep 1827Order Schizaeales Schimp Traiteacute Paleacuteont Veacuteg 1 674 Mar 1869Subclass Schizaeidae Doweld New Syllabus 346 May 2006Suborder Schizaeineae Bessey Bot High Schools 377 14ndash21 Aug 1880 (ldquoSchizaeaceaerdquo)Class Schizaeopsida Doweld Tent Syst Pl Vasc x 23 Dec 2001bullFam Sinopteridaceae Koidz Acta Phytotax Geobot 3 50 30 Mayndash20 Jul 1934 bullFam Sphaerostephanaceae Ching Sunyatsenia 5 240 1940 nom nud without Latin descriptionbullFam Stenochlaenaceae Ching Acta Phytotax Sin 16(4) 18 Nov 1978bullFam Stromatopteridaceae Bierh Phytomorphology 18 263 15 Dec 1968 ndash basionym Stromatopteridoideae Nakai Bull Natl Sci

Mus Tokyo 29 32 1950Order Stromatopteridales Pic Serm ex Reveal Phytologia 74 176 25 Mar 1993bullFam Taenitidaceae Pic Serm Webbia 29 1 10 Feb 1975bullFam Tectariaceae Panigrahi J Orissa Bot Soc 8 41 1986bullFam Thelypteridaceae Pic Serm Webbia 24 709 28 Apr 1970Order Thelypteridales Doweld Tent Syst Pl Vasc xi 23 Dec 2001bullFam Thyrsopteridaceae C Presl Gefaumlssbuumlndel Farrn 22 38 1847 (ldquoThyrsopterideaerdquo) nom rej in favor of Dicksoniaceae (1848)bullFam Tmesipteridaceae Nakai Chosakuronbun Mokuroku [Ord Fam Trib Nov] 206 1943bullFam Trichomanaceae Burmeist Handb Naturgesch 1 196 12ndash17 Dec 1836 (ldquoTrichomanoideaerdquo)bullFam Vittariaceae Ching Sunyatsenia 5 210 232 30 Oct 1940 ndash basionym Vittarieae C Presl [Tent Pterid] Abh Koumlnigl Boumlhm

Ges Wiss ser 4 5 164 before 2 Dec 1836 (ldquoVittariaceaerdquo)bullFam Woodsiaceae Herter Revista Sudamer Bot 9 14 Jun 1949 ndash basionym Woodsieae A Gray Man Bot ed 2 588 1 Sep 1856


731

Appendix 5 ContinuedDrynaria (37) Leucostegia (32) Ophioderma = Ophioglossum Pyrrosia (37)Dryoathyrium = Deparia Lindsaea (23) Ophioglossum (1) Quercifilix = TectariaDryopolystichum (32) Lindsayoides = Nephrolepis Oreopteris = Thelypteris Radiovittaria (26)Dryopsis (32) Lithostegia (32) Ormoloma (23) Regnellidium (13)Dryopteris (32) Litobrochia = Pteris Orthiopteris = Saccoloma Revwattsia (32)Edanyoa = Bolbitis Llavea (26) Osmunda (5) Rhachidosorus (29)Egenolfia = Bolbitis Lomagramma (32) Pachypleuria (36) Rheopteris (26)Elaphoglossum (32) Lomaphlebia = Grammitis Paesia (25) Rosenstockia = HymenophyllumEnterosora (37) Lomaria = Blechnum Paltonium = Neurodium Rumohra (32)Equisetum (3) Lomariopsis (33) Papuapteris = Polystichum Saccoloma (24)Eriosorus (26) Lonchitis (23) Paraceterach (26) Sadleria (30)Fadyenia = Tectaria Lophosoria (21) Parahemionitis (26) Saffordia = TrachypterisFeea = Trichomanes Lorinseria = Woodwardia Parasorus = Davallia Sagenia = TectariaFourniera = Sphaeropteris Loxogramme (37) Parathelypteris = Thelypteris Salpichlaena (30)Glaphyropteridopsis = Cyclosorus Loxoma (16) Pecluma (37) Salvinia (14)Glaphyropteris = Cyclosorus Loxoscaphe = Asplenium Pelazoneuron = Cyclosorus Sceptridium = BotrychiumGleichenella (7) Loxsomopsis (16) Pellaea (26) Schaffneria = AspleniumGleichenia (7) Luisma (37) Peltapteris = Elaphoglossum Schizaea (12)Goniophlebium (37) Lunathyrium = Deparia Pentagramma (26) Scleroglossum (26)Goniopteris = Cyclosorus Lygodium (10) Pentarhizidium (31) Scoliosorus (26)Gonocormus = Crepidomanes Macroglena = Abrodictyum Peranema (32) Scyphularia = DavalliaGrammitis (37) Macrothelypteris (28) Phanerophlebia (32) Selliguea (37)Gymnocarpium (29) Mankyua (1) Phanerosorus (9) Serpocaulon (37)Gymnogramma = Hemionitis Marattia (4) Phegopteris (28) Serpyllopsis = HymenophyllumGymnogrammitis (37) Marginariopsis = Pleopeltis Phlebodium (37) Sinephropteris = AspleniumGymnopteris = Hemionitis Marsilea (13) Photinopteris = Aglaomorpha Sinopteris = AleuritopterisGymnosphaera = Alsophila Matonia (9) Phyllitis = Asplenium Solanopteris = MicrogrammaHaplopteris (26) Matteuccia (31) Phymatosorus (37) Sphaerocionium =Hecistopteris (26) Maxonia (32) Pilularia (13) HymenophyllumHelminthostachys (1) Mecodium = Hymenophyllum Pityrogramma (26) Sphaeropteris (20)Hemidictyum (29) Megalastrum (32) Plagiogyria (18) Sphaerostephanos = CyclosorusHemigramma = Tectaria Melpomene (37) Platycerium (37) Sphenomeris (23)Hemionitis (26) Meniscium = Cyclosorus Platygyria = Lepisorus Steenisioblechnum (30)Hemitelia = Cyathea Menisorus = Cyclosorus Platyloma (26) Stegnogramma = CyclosorusHeterogonium (34) Merinthosorus = Aglaomorpha Platyzoma (26) Steiropteris = CyclosorusHippochaete = Equisetum Meryngium = Hymenophyllum Plecosorus = Polystichum Stenochlaena (30)Histiopteris (25) Mesophlebion = Cyclosorus Pleocnemia (34) Stenolepia (32)Holcochlaena (26) Metathelypteris = Thelypteris Pleopeltis (37) Sticherus (7)Holodictyum = Asplenium Metaxya (22) Plesioneuron = Cyclosorus Stigmatopteris (32)Holostachyum = Aglaomorpha Microgonium = Didymoglossum Pleuroderris = Tectaria Stromatopteris (7)Homalosorus (29) Microgramma (37) Pleurosoriopsis (37) Synammia (37)Humata = Davallia Microlepia (25) Pleurosorus = Asplenium Syngramma (26)Hyalotricha = Campyloneurum Micropolypodium (37) Pneumatopteris = Cyclosorus Taenitis (26)Hyalotrichopteris = Campyloneurum Microsorum (37) Podosorus (37) Tapeinidium (23)Hymenasplenium (27) Microstaphyla = Elaphoglossum Polybotrya (32) Tectaria (34)Hymenocystis = Woodsia Microtrichomanes = Polyphlebium (6) Teratophyllum (32)Hymenoglossum = Hymenophyllum Hymenophyllum Polypodioides (37) Terpsichore (37)Hymenophyllopsis (20) Mildella (26) Polypodiopteris (37) Thamnopteris = AspleniumHymenophyllum (6) Mohria = Anemia Polypodium (37) Thelypteris (28)Hypodematium (32) Monachosorum (25) Polystichopsis (32) Themelium (37)Hypoderris (34) Monogramma (26) Polystichum (32) Thylacopteris (37)Hypolepis (25) Monomelangium = Diplazium Polytaenium (26) Thyrsopteris (15)Idiopteris = Pteris Neocheiropteris (37) Pronephrium = Cyclosorus Thysanosoria (33)Jamesonia (26) Nephelea = Alsophila Prosaptia (37) Tmesipteris (2)Japanobotrychium = Botrychium Nephopteris (26) Protowoodsia (29) Todea (5)Kontumia (37) Nephrolepis (33) Psammiosorus (34) Trachypteris (26)Kuniwatsukia = Athyrium Neurocallis (26) Pseudocolysis = Pleopeltis Trichoneuron (32)Lacostea = Trichomanes Neurodium (37) Pseudocyclosorus = Cyclosorus Trichipteris = CyatheaLacosteopsis = Vandenboschia Neuromanes = Trichomanes Pseudocystopteris (29) Trichomanes (6)Lastrea = Thelypteris Niphidium (37) Pseudodrynaria = Aglaomorpha Trigonospora = CyclosorusLastreopsis (32) Notholaena (26) Pseudophegopteris (28) Triplophyllum (34)Lecanium = Didymoglossum Nothoperanema = Dryopteris Pseudotectaria = Tectaria Trismeria = PityrogrammaLecanopteris (37) Ochropteris (26) Psilotum (2) Vaginularia = MonogrammaLellingeria (37) Odontosoria (23) Psomiocarpa (34) Vandenboschia (6)Lemmaphyllum (37) Oenotrichia (25) Pteridium (25) Vittaria (26)Lepisorus (37) Oenotrichia pp (32) Pteridoblechnum (30) Weatherbya = LemmaphyllumLeptochilus (37) Oleandra (35) Pteridrys (34) Woodsia (29)Leptogramma = Cyclosorus Olfersia (32) Pteris (26) Woodwardia (30)Leptolepia (25) Onoclea (31) Pterozonium (26) Xiphopteris = CochlidiumLeptopteris (5) Onocleopsis (31) Ptilopteris = Monachorosum Xyropteris (23)Leptorumohra = Arachniodes Onychium (26) Pycnodoria = Pteris Zygophlebia (37)

lizing data from morphology seven chloroplast markers(rbcL atpA atpB accD rps4 16S rDNA ITS) one nu-clear gene (18S rDNA) and three mitochondrial genes(atp1 nad2 nad5) prompt us to reevaluate the classifi-cation of these vascular plants Multiple-gene phyloge-netic analyses eg studies by Wolf (1996) Wolf amp al(1998) Pryer amp al (2001a 2004b) Schneider amp al(2004c) Wikstroumlm amp Pryer (2005) and Schuettpelz amp al(in press) have given rise to growing conviction in boththe composition and relationships of taxa at familial andordinal ranks Five recent morphological analyses ofrelationships by Pryer amp al (1995 2001a) Schneider(1996a) Stevenson amp Loconte (1996) and Schneider ampal (in prep) have increased support for the molecular-based consensus topology For eusporangiate and basalleptosporangiate ferns evidence is now sufficient to al-low us to circumscribe confidently most clades and as-sign ranks However for some more derived leptospo-rangiate ferns the phylogenetic evidence is still some-what equivocal Consequently declaration of phyloge-netic positions for some taxa and the assignment of ranks(which we consider subjective and secondary) in a clas-sical ldquoLinnaean-stylerdquo hierarchy are tentative In this pa-per we present a revised view of the classification of ex-tant ferns taking into account all relevant evidence Wefocus our classification at the ranks of class order andfamily believing that the information at hand is mostappropriate for resolution and understanding of relation-ships at these levels Within most families and especial-ly at the generic level there is still insufficient evidenceto attempt many classificatory decisions

In the classification proposed herein we account forand place in a revised taxonomic framework all names atfamily and ordinal rank utilized in previous major classifi-cations directed at ferns particularly those that have beenproposed in the last eighty years Bower (1926) Christen-sen (1938) Ching (1940 1978) Dickason (1946) Cope-land (1947) Holttum (1947 1949 1973) Pichi Sermolli(1958 1977) Mehra (1961) Wagner (1969) Nayar(1970) Tagawa amp Iwatsuki (1972) Mickel (1974) Try-on amp Tryon (1982) Kramer (in Kubitzki 1990) Hennip-man (1996) and Stevenson amp Loconte (1996) We alsoconsider various herbarium schemes in common usesuch as the one by Crabbe amp al (1975) Many of theseclassifications as well as others have been reviewed andreproduced by Pichi Sermolli (1973) who also summa-rized detailed information on family names of ferns(1970 1981 1982 1986) Smith (1995) provided a re-cent summary and discussion of these classifications

Classifications serve many purposes among them toprovide a genealogical framework in which to identifyplants organize herbaria retrieve information and toconduct many kinds of studies (eg evolutionary mor-phological and physiological) Multi-use classifications

serve us best if we name only those clades that are read-ily recognizable and characterized by morphologicalsynapomorphic characters at least at family and higherranks However more traditional (morphology-based)and practical classifications are sometimes incompatiblewith the results and classifications implied by phyloge-netic studies especially when the principle of monophy-ly is used as a grouping criterion (recognizing clades andnot paraphyletic grades APG II 2003) When a tradi-tionally recognized family nests within another complexclassificatory choices ensue (1) recognition of para-phyletic families (Brummitt 1996 1997 Lideacuten amp al1997 Moore 1998 Diggs amp Lipscomb 2002 Grant2003) (2) dismemberment of a recognized family intosmaller families (eg the disintegration of classicalScrophulariaceae Olmstead amp al 2001) or (3) integra-tion of the traditional family that causes the paraphylyinto the ldquoprogenitorrdquo family The first choice preferredby some leads to recognition of unnatural (non-mono-phyletic) groups which in our opinion often retards orobscures investigation into interesting biological phyto-geographic and evolutionary questions The second so-lution supposes that we have morphological synapomor-phies for nodes that lead to all of the segregate familiesand this is not often the case although one hopes thateventually we shall find these synapomorphies Untilthen it may be nearly impossible to define some segre-gate families in such a way that they would be bothkeyable and circumscribable The third solution is a ldquofastfixrdquo to the problem but expediency often demands thatat least some of the intrafamilial subclades also be rec-ognized taxonomically either at a lower rank (eg sub-family) or with an unranked informal name (eg ldquogram-mitid fernsrdquo) until further decisions on rank can be madeThis third option seems to us the most practicable andpractical solution toward a ldquofirst-passrdquo revision of fernclassification As more data are gathered and future phy-logenetic analyses provide better resolved and better sup-ported topologies one expects further insight into identi-fying synapomorphies for segregate taxa enabling even-tual movement toward the second option of recognizingsegregate monophyletic families

The nested linearity of a ranked Linnaean classifica-tion and the availability of a limited number of univer-sally accepted ranks facilitate the organization and ar-rangement of taxa for cataloging purposes such as theefficient arrangement of specimens in herbaria or theorganization and retrieval of regional biodiversity fromchecklists floras and other taxonomic products Rankedclassifications often perform poorly however whencalled upon to present explicit statements of clade mem-bership and relationship Such classifications also canmislead one into making specious comparisons eg at-tempting to describe the extent of biodiversity in terms of


706











707








Polypodio-psida










Polypodiales




Dryopteridaceae


Leptosporangiates


Ferns

Polypods

Eupolypods

Eupolypods II

Eupolypods I


Culcitaceae


708















709












710












711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731












707








Polypodio-psida










Polypodiales




Dryopteridaceae


Leptosporangiates


Ferns

Polypods

Eupolypods

Eupolypods II

Eupolypods I


Culcitaceae


708















709












710












711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731









Polypodio-psida










Polypodiales




Dryopteridaceae


Leptosporangiates


Ferns

Polypods

Eupolypods

Eupolypods II

Eupolypods I


Culcitaceae


708















709












710












711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731
















709












710












711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731













710












711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731













711












712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731













712









713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731










713









714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731










714











715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731












715









716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731










716









717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731










717










718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731











718










719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731











719



































720






































721










































722
































723









































724






725



726







727









728












729






730






731




































720






































721










































722
































723









































724






725



726







727









728












729






730






731







































721










































722
































723









































724






725



726







727









728












729






730






731











































722
































723









































724






725



726







727









728












729






730






731

































723









































724






725



726







727









728












729






730






731










































724






725



726







727









728












729






730






731







725



726







727









728












729






730






731



726







727









728












729






730






731



727









728












729






730






731



728












729






730






731



729






730






731



730






731



731


Aclassification for extant ferns - Sites@Duke · leptosporangiate ferns, evidence is now sufficient...

Documents

Transcript of Aclassification for extant ferns - Sites@Duke · leptosporangiate ferns, evidence is now sufficient...