A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data

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1 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data Eric Schuettpelz, Cheng-Wei Chen,2 Michael Kessler, Jerald B. Pinson,4 Gabriel Johnson, lex Davila,5 lyssa T. Cochran,6 Layne uiet7 & Kathleen M. Pryer7 Department of Botany, National Museum of Natural istory, Smithsonian Institution, Washington, D.C. 200, U.S.. 2 Division of Botanical Garden, Taiwan Forestry Research Institute, Taipei 0066, Taiwan Institute of Systematic Botany, University of Zurich, 8008 Zurich, Switzerland 4 Department of Biology, University of Florida, Gainesville, Florida 26, U.S.. 5 Department of Biology and Marine Biology, University of North Carolina Wilmington, Wilmington, North Carolina 2840, U.S.. 6 Department of Biology, Colorado State University, Fort Collins, Colorado 8052, U.S.. 7 Department of Biology, Duke University, Durham, North Carolina 27708, U.S.. uthor for correspondence: Eric Schuettpelz, schuettpelze@si.edu ORICD ES, DOI bstract ittarioid ferns compose a well-supported clade of 00 0 species of highly simplified epiphytes in the family Pteridaceae. Generic circumscriptions within the vittarioid clade were among the first in ferns to be evaluated and revised based on molecular phylogenetic data. Initial analyses of rbcl sequences revealed strong geographic structure and demonstrated that the two largest vittarioid genera, as then defined, each had phylogenetically distinct merican and Old World components. The results of subsequent studies that included as many as 6 individuals of species, but still relied on a single gene, were generally consistent with the early findings. ere, we build upon the previous datasets, incorporating many more samples (8 individuals representing 72 species) and additional plastid markers (atp, chln, rbcl, rpo). nalysis of our larger dataset serves to better characterize known lineages, reveals new lineages, and ultimately uncovers an underlying geographic signal that is even stronger than was previously appreciated. In our revised generic classification, we recognize a total of eleven vittarioid genera. Each genus, including the new genus ntrophyopsis (Benedict) Schuettp., stat. nov., is readily diagnosable based on morphology, with micromorphological characters related to soral paraphyses and spores complementing more obvious features such as venation and the distribution of sporangia. key to the currently recognized vittarioid genera, brief generic descriptions, and five new species combinations are provided. Keywords frica; epiphytes; paraphyses; phylogeny; pteridophytes; spores; taxonomic revision Supplementary Material Electronic Supplement (Fig. S) and DN sequence alignments are available in the Supplementary Data section of the online version of this article at INTRODUCTION In ferns, as in most branches of the tree of life, phylogenetic analyses of molecular data have greatly improved our ability to identify natural groupings that are subsequently reflected in classifications grounded in the principle of monophyly (Smith & al., 2006; Rothfels & al., 202; Christenhusz & Chase, 204). In some cases, the results of such analyses are consistent with earlier notions of relationships inferred from morphological features (Schneider & al., 2009). owever, in many other instances, lineages are revealed that are morphologically confounding and we struggle to identify synapomorphies (Sundue & Rothfels, 204). The pursuit of such defining characteristics is especially problematic when working within a group possessing very limited morphological disparity. The well-defined vittarioid fern clade consists of 00 0 (Lindsay, 200) highly simplified and predominantly epiphytic species (Fig. ). These plants, characterized by the presence of silica bodies (Sundue, 2009) but a lack of sclerenchyma (Bower 928; Ruhfel & al., 2008), were long regarded as composing a distinct family ittariaceae (Ching, 940; Tryon & Tryon, 982; Kramer, 990). owever, phylogenetic analyses have demonstrated that these ferns nest well within the Pteridaceae (Crane & al., 995; asebe & al., 995; Prado & al., 2007; Schuettpelz & al., 2007), as sister to the genus diantum L. (Lu & al., 202; Rothfels & Schuettpelz, 204; Pryer & al., 206). The vittarioids have been variously partitioned through time (Benedict, 9; Williams, 927; Copeland, 947). In the years leading up to the first molecular phylogenetic analyses of these ferns, six genera were commonly recognized based primarily on leaf division, venation, and the distribution of Received: 7 Dec 205 returned for (first) revision: Mar 206 (last) revision received: 9 May 206 accepted: 9 May 206 publication date(s): online fast track, n/a; in print and online issues, 0 ug 206 International ssociation for Plant Taxonomy (IPT) ersion of Record

2 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns Fig.. ittarioid fern habits and key morphological features., Epiphytic habit of aplopteris elongata (Sw.) E..Crane (leaves ca. cm wide); B, Epiphytic habit, and forked leaves, of ecistopteris pumila (Spreng.) J.Sm. (leaves ca. cm long); C, Epiphytic habit of ntrophyum annamense Tardieu & C.Chr. (leaves ca. 5 cm wide); D, Epiphytic habit of aplopteris graminea (Poir.) comb. ined. (= Monogramma graminea (Poir.) Schkuhr, leaves ca. 5 cm long): E, Pinnately compound leaves and round sori of Rheopteris cheesmaniae lston (leaves ca. cm wide); F, Biseriate areolate venation of aplopteris elongata (leaf ca. cm wide); G, Pluriseriate areolate venation of ntrophyum sessilifolium (Cav.) Spreng. (leaf about cm wide);, Sori of Radiovittaria remota (Fée) E..Crane, restricted to submarginal commissures (leaf ca. cm wide); I, Sori of ntrophyum castaneum.ito, along reticulate veins (leaf ca. cm wide); J, Sori of Scoliosorus ensiformis (ook.) T.Moore, along reticulate veins (leaf ca. 2 cm wide); K, Sori of aginularia acrocarpa olttum, appearing as one continuous line on each leaf (leaves ca. mm wide); L, Soral paraphyses of ntrophyum callifolium Blume, with slender apical cells (scale bar 00 µm); M, Soral paraphyses of ntrophyum subfalcatum Brack., with spherical apical cells (scale bar 00 µm); N, Soral paraphyses of aplopteris elongata, with obconic apical cells (scale bar 00 µm). Image credits: Cheng-Wei Chen (C, G, I, K, L, M, N); Fay-Wei Li (B); Germinal Rouhan (D); Eric Schuettpelz (, F); Michael Sundue, (, J); and Wayne Takeuchi (E). ersion of Record 709

3 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: sporangia (Tryon & Tryon, 982; Kramer, 990). The monotypic Rheopteris lston was considered to be unique (among vittarioids) based on its pinnately divided leaves and round sori (Fig. E). The small genus ecistopteris J.Sm. was also easily segregated based on its minute, forked or lobed leaves (Fig. B). The remaining genera were characterized by the presence of simple leaves. mong them, the monotypic netium Splitg. was readily separated from other vittarioids by having sporangia on its abaxial leaf surfaces that are not restricted to the veins. ittaria Sm., with species recognized at the time (Kramer, 990), was defined as having sporangia confined to submarginal commissures (Fig. ). ntrophyum Kaulf. was applied to those species (about 50) with pluriseriate areolate venation (Fig. G) and sporangia following the veins (not confined to submarginal commissures; Fig. I). Finally, Monogramma Comm. ex Schkuhr consisted of a handful of diminutive species with sporangia borne along the singular leaf vein, or the lateral vein(s), if present (Fig. K). Generic circumscriptions within vittarioids were among the first in ferns to be evaluated and revised based on molecular phylogenetic data. lthough initial analyses of rbcl sequences (Crane & al., 995) included only species of four of the six genera widely recognized at the time (Rheopteris and Monogramma were not sampled), they already revealed a significant lack of monophyly at the generic level. Specifically, ittaria was demonstrated to include four distinct elements and ntrophyum three. new phylogenetically based, and morphologically defensible, classification was thus proposed that divided ittaria and ntrophyum (sensu Tryon & Tryon, 982; Kramer, 990) into four genera (nanthacorus Underw. & Maxon, aplopteris C.Presl, Radiovittaria (Benedict) E..Crane, ittaria) and three genera (ntrophyum, Polytaenium Desv., Scoliosorus T.Moore), respectively (Crane, 997). Names existed at the generic level for each of these segregates, with one exception for which a subgeneric name was elevated (Radiovittaria). Crane s (997) classification relied heavily on micromorphological characters, focusing specifically on spore type and the shape of the apical cells of the soral paraphyses (Fig. L N), but also considered gross morphology and geography. Of the species with sporangia confined to submarginal commissures (formerly placed in ittaria), nanthacorus was readily segregated based on its pluriseriate areolate venation (all other elements had biseriate areolate venation), ittaria was restricted to taxa with soral paraphyses having slender apical cells, and species with obconic apical cells were further divided based on their phyllotaxy (distichous in aplopteris and polystichous in Radiovittaria). Of the species with pluriseriate areolate venation and sporangia following the veins (not confined to submarginal commissures; formerly placed in ntrophyum), those lacking soral paraphyses were placed in Polytaenium and those having paraphyses were placed in ntrophyum or Scoliosorus (depending on whether their spores were trilete or monolete, respectively). Subsequent phylogenetic studies (Schuettpelz & al., 2007; Ruhfel & al., 2008) included the previously unsampled genera Rheopteris and Monogramma. The monotypic Rheopteris and some species of Monogramma (as then defined) were together found to be sister to the remaining vittarioids; other species of Monogramma were resolved elsewhere within the vittarioid phylogeny. Ruhfel & al. (2008) advocated for the application of the existing name aginularia Fée to those species formerly treated in Monogramma that were most closely related to Rheopteris. Lindsay & Chen (204) placed another species formerly treated in Monogramma in aplopteris, noting that the type species of Monogramma had other affinities. Recent analyses focused attention on relationships within the two largest vittarioid genera aplopteris and ntrophyum (Chen & al., 20a, b, 205). owever, a well-sampled study of the overall vittarioid phylogeny, based on multiple molecular markers, has yet to be published. ere, we build upon the earlier analyses of vittarioid fern relationships by incorporating many more samples and additional plastid markers. nalysis of our larger dataset serves to better characterize known lineages, reveals new lineages, and ultimately uncovers an underlying geographic signal that is even stronger than was previously recognized. MTERILS ND METODS The ingroup sampling for this study comprised 8 individuals of 72 vittarioid species (ppendix ). n effort was made to include as many species as possible, but also to sample from across the ranges of the more widespread species. In general, no more than one individual of a given species was sampled from a particular country. owever, if representatives were only obtainable from a single country, two individuals were included from that country (as available). In addition to the 8 vittarioid samples, 7 individuals representing 7 species from across diantum (ppendix ) were included to root the vittarioid phylogeny, based on earlier studies that found a sister relationship between diantum and the vittarioids (Schuettpelz & al., 2007; Lu & al., 202; Rothfels & Schuettpelz, 204; Pryer & al., 206). Genomic DN was typically extracted from silica-dried samples (most individuals) or herbarium fragments using a modified CTB protocol (Doyle & Doyle, 987), as described in detail in Beck & al. (20). Four plastid gene regions (atp, chln, rbcl, rpo) were amplified independently using the polymerase chain reaction (PCR). Nuclear gene sequencing within ferns is challenging (Schuettpelz & al., 2008) and nuclear markers were not included here. owever, with the greater availability of transcriptome data and next-generation sequencing approaches, the landscape is changing (Rothfels & al., 205). For atp, either a large region (including the entire atp gene, plus portions of the atpf and trnr genes and the associated spacers) or a small region (including only a portion of the atp gene) was targeted, depending mostly on the quality of each genomic DN extraction. For the larger region, the primers (Table ) and amplification protocol of Schuettpelz & al. (2006) were utilized, whereas for the smaller region, the primers (Table ) and amplification protocol of Cochran & 70 ersion of Record

4 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns Table. mplification and sequencing primers used in this study of vittarioid ferns. Region Primer Utility (direction) Reference or sequence (if new) atp (large) ESTPF42F mplification /sequencing (forward) Schuettpelz & al., 2006 atp (large) ESTP55F Sequencing (forward) Schuettpelz & al., 2006 atp (large) ESTP557R Sequencing (reverse) Schuettpelz & al., 2006 atp (large) ESTP856F Sequencing (forward) Schuettpelz & al., 2006 atp (large) ESTP877R Sequencing (reverse) Schuettpelz & al., 2006 atp (large) ESTRNR46F mplification /sequencing (reverse) Schuettpelz & al., 2006 atp (small) atp-f mplification /sequencing (forward) Cochran & al., 204 atp (small) atp-r mplification /sequencing (reverse) Cochran & al., 204 chln chln-f2 mplification /sequencing (forward) CGWTYGCRYGGCGTYGSG chln chln-r2 mplification /sequencing (reverse) CWTTTTTTCGTCCRGCRCGTG rbcl ESRBCLF mplification /sequencing (forward) Schuettpelz & Pryer, 2007 rbcl ESRBCL628F Sequencing (forward) Schuettpelz & Pryer, 2007 rbcl ESRBCL654R Sequencing (reverse) Schuettpelz & Pryer, 2007 rbcl ESRBCL6R mplification /sequencing (reverse) Schuettpelz & Pryer, 2007 rpo rpo-f mplification /sequencing (forward) TRCYGGTTTCYCTCGGG rpo rpo-r mplification /sequencing (reverse) TTRGCTCTRGCRGGTRTTC al. (204) were applied. For chln (a plastid gene coding for photochlorophyllide reductase subunit N), primers were newly developed (Table ), but the amplification protocol of Cochran & al. (204) was followed. For rbcl, a large segment of the gene was generally amplified (following Schuettpelz & Pryer, 2007); however, in some cases it was necessary to amplify this segment in two parts due to the quality of the genomic DN extraction. For rpo (a plastid gene coding for the RN polymerase alpha subunit), primers were again newly developed (Table ) and the protocol of Cochran & al. (204) was followed, except that the annealing temperature was reduced to 45 C. isualization and cleanup of PCR products, as well as sequencing, followed previously published protocols (Schuettpelz & al., 2006; Schuettpelz & Pryer, 2007; Cochran & al., 204). For some individuals, sequences were already published and a total of 0 sequences were thus simply downloaded from GenBank. Because of failed amplification or sequencing reactions, 55 sequences could not be obtained, but a total of 45 sequences were newly acquired for this study and uploaded to GenBank (ppendix ). For each plastid gene region, the relevant sequences were assembled in liiew v..7 (Larsson, 204) and initially aligned therein with MUSCLE v..8 (Edgar, 2004). The resulting alignments were then visually inspected, and manually edited with the guidance of corresponding amino acid translations. Because the atp sequences obtained were of two distinct lengths (resulting in copious amounts of missing data in the alignment for some taxa) and also because the longer sequences incorporated non-coding flanking regions (where alignment was frequently ambiguous), the characters outside of the shorter targeted region were excluded. Details relevant to the datasets are provided in Table 2. The four single-gene alignments were independently subjected to phylogenetic analysis in MrBayes v..2. (uelsenbeck & Ronquist, 200; Ronquist & uelsenbeck, 200) using the GTR + G model of sequence evolution and a temperature of nalyses incorporated four independent runs, each with four chains, proceeded for 0 million generations, and sampled trees every 8000 generations. fter completion, output files were examined for reasonable chain swap rates. To Table 2. Details concerning the alignments used in this study of vittarioid ferns. Dataset Individuals Total characters Included characters ariable characters Missing data (%)* atp chln rbcl rpo Combined *Calculation based on included characters. ersion of Record 7

5 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: verify that the runs had converged, the standard deviation of split frequencies was evaluated; additionally, parameter estimates were plotted and ESS values examined in Tracer v..6 (Rambaut & al., 204). Based on these diagnostics, the first 250 trees (corresponding to the first 2 million generations) were very conservatively excluded before obtaining a majority-rule consensus phylogeny with clade posterior probabilities for each plastid gene region. The four single-gene trees were rooted at the split between diantum and vittarioids and inspected visually for well-supported (posterior probability, PP 0.95) topological differences. Only three significant topological differences were encountered among the four gene trees, all involving shallow branches. While atp supported the monophyly of the two included samples of aplopteris humblotii (ieron.) S.Linds. & C.W.Chen (945, 946), rbcl found support for their paraphyly relative to. guineensis (Desv.) E..Crane and. schliebenii (Reimers) Schuettp. Likewise, while chln supported the monophyly of the two included samples of ittaria isoetifolia Bory (942, 06), rpo found support for their paraphyly relative to a sample of. lineata (L.) Sm. (8927). Finally, while chln supported the two included samples of aplopteris mediosora (ayata) X.C.Zhang (9694, 9695) as sister to a clade comprising. doniana (Mett. ex ieron.) E..Crane,. fudzinoi (Makino) E..Crane,. linearifolia (Ching) X.C.Zhang,. plurisulcata (Ching) X.C.Zhang, and. taeniophylla (Copel.) E..Crane, rbcl supported them as sister to. malayensis (olttum) E..Crane. Because the aforementioned conflicts were all inconsequential to the primary objectives of this study, the atp, chln, rbcl, and rpo datasets were combined for further analysis. The combined dataset was phylogenetically analyzed using a maximum likelihood approach, as implemented in RxML v.8.0. (Stamatakis, 204). This analysis employed the GTRGMM model of sequence evolution, with parameters independently estimated for each gene, and involved 000 rapid bootstrap inferences followed by a thorough maximum likelihood search. The combined dataset was also analyzed using the Bayesian approach described above for the single-gene alignments, but with model parameters estimated separately for each gene and each run proceeding for 40 million generations. ere, trees were sampled every 2,000 generations and convergence was assessed as above; the first 8 million generations were excluded prior to calculating the majority-rule consensus phylogeny with clade posterior probabilities. For each vittarioid species in our dataset, we assessed leaf venation pattern, the apical cell shape of soral paraphyses, spore type, geographic distribution, and, as applicable, other morphological features by examining herbarium specimens and the relevant literature. RESULTS The monophyly of the vittarioids relative to diantum is strongly supported, with a maximum likelihood bootstrap percentage (BS) of 00 and a Bayesian posterior probability (PP) of.00 (node, Fig. 2). Within vittarioids, the first divergence is highly asymmetrical with regard to species number. The smaller clade (BS = 00; PP =.00; node 2, Figs. 2, ) is characterized by free venation (pinnately compound or indistinct pinnate) and trilete spores. It consists exclusively of species from sia and the Pacific Ocean and unites the only vittarioid genus with compound leaves (Rheopteris) with some of the most morphologically simplified species (treated here in aginularia). The larger vittarioid clade (BS = 00; PP =.00; node, Figs. 2, ) consists primarily of simple-leaved species with reticulate venation but is otherwise morphologically variable. Within this larger clade, a rather symmetrical divergence separates two subclades of approximately equal size. One subclade (BS = 00; PP =.00; node 4, Figs. 2, ) is characterized by paraphyses with obconic apical cells and consists almost entirely of species with biseriate areolate leaf venation. The other subclade (BS = 98; PP =.00; node 5, Figs. 2, ) does not have an obvious unifying morphological feature, but includes all of the species with pluriseriate areolate leaf venation (along with six sampled species possessing biseriate areolate venation and one with parallel free veins). Within the two aforementioned subclades, a consistent biogeographic pattern emerges, with each containing a well-supported merican lineage sister to a well-supported Old World lineage. In each of the two Old World lineages, there is, in turn, an frican / Indian Ocean group sister to a largely sian / Pacific Ocean group (the latter with one or a few derived frican / Indian Ocean representatives). Overall, relationships within the vittarioids are reasonably well supported. Of 6 resolved nodes, 06 received good support (i.e., BS 70 and PP 0.95; Electr. Suppl.: Fig. S). Nearly all (29 of 0) poorly supported nodes involve relationships among species within genera. In most (8 of 42) cases where more than one sample of a species was available for sequencing, species were resolved as monophyletic (6 of 42 received good support; Electr. Suppl.: Fig. S). DISCUSSION Phylogeny. The earliest molecular phylogenetic study of vittarioid ferns (Crane & al., 995) found the two largest genera, as commonly circumscribed at the time (ntrophyum, ittaria), to be polyphyletic. This motivated a revised classification for the group (Crane, 997) that was able to accommodate the goal of only recognizing monophyletic genera simply through the Fig. 2. ittarioid fern phylogeny resulting from maximum likelihood analysis of a four-gene (atp, chln, rbcl, rpo) plastid dataset incorporating 8 ingroup and 7 outgroup individuals. Maximum likelihood bootstrap percentages (BS) and Bayesian posterior probabilities (PP) are provided at select nodes (BS / PP; C = 00; I =.00; < = BS < 50 or PP < 0.50); support values for all nodes are provided in Fig. S (Electr. Suppl.). White numbers in black circles at nodes correspond to major clades / subclades discussed in the text. Species names in tree follow the generic classification proposed in this study; for previous generic assignments, see Fig.. Numbers following species names are Fern Lab Database ( biology.duke.edu) voucher numbers (ppendix ). 72 ersion of Record

6 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns 0.05 substitutions per site 8/I diantum capillus-veneris 4609 diantum malesianum 2506 diantum tenerum 2504 diantum formosum 4602 diantum andicola 5549 diantum tetraphyllum 2505 diantum peruvianum 2507 Rheopteris cheesmaniae 7 aginularia acrocarpa aginularia acrocarpa 900 aginularia acrocarpa 75 aginularia trichoidea 4668 aginularia trichoidea 909 aginularia angustissima 9705 aginularia paradoxa 906 aginularia paradoxa 9699 ecistopteris pumila 55 ntrophyopsis manniana 9702 ecistopteris pumila 278 ntrophyopsis manniana 940 Radiovittaria ruiziana 977 ntrophyopsis bivittata 996 Radiovittaria ruiziana 979 ntrophyopsis boryana 9076 Radiovittaria moritziana 9046 ntrophyopsis boryana 970 Radiovittaria moritziana 9059 ntrophyum obovatum 842 Radiovittaria latifolia 9058 ntrophyum castaneum 9675 Radiovittaria latifolia 9052 ntrophyum latifolium 078 Radiovittaria minima 855 ntrophyum subfalcatum /I Radiovittaria gardneriana 445 ntrophyum plantagineum 9680 Radiovittaria gardneriana 870 ntrophyum parvulum 9679 Radiovittaria gardneriana 905 ntrophyum parvulum 900 Radiovittaria remota 854 ntrophyum semicostatum 9066 Radiovittaria stipitata 55 66/0.90 ntrophyum semicostatum 845 Radiovittaria stipitata 249 ntrophyum formosanum Radiovittaria stipitata 9045 ntrophyum henryi 9677 aplopteris graminea 058 ntrophyum coriaceum 99 aplopteris graminea 548 ntrophyum annamense 967 aplopteris volkensii 94 ntrophyum sessilifolium 8999 aplopteris humblotii 946 ntrophyum sessilifolium 968 aplopteris humblotii 945 ntrophyum malgassicum 998 aplopteris schliebenii 9407 ntrophyum vittarioides 968 aplopteris guineensis 960 ntrophyum wallichianum /I 98/I aplopteris guineensis ntrophyum wallichianum 995 aplopteris heterophylla 969 ntrophyum reticulatum 844 aplopteris flexuosa 9688 ntrophyum callifolium 908 aplopteris flexuosa 4786 aplopteris amboinensis 97 ntrophyum callifolium 9674 ntrophyum callifolium 99 aplopteris amboinensis 9684 ntrophyum callifolium 408 aplopteris scolopendrina Polytaenium brasilianum aplopteris scolopendrina 905 Polytaenium brasilianum 9087 aplopteris scolopendrina 906 Polytaenium citrifolium 906 aplopteris malayensis 44 Polytaenium citrifolium 9 aplopteris mediosora 9694 Polytaenium citrifolium 5 aplopteris mediosora 9695 Polytaenium lineatum 9062 aplopteris himalayensis /0.7 Polytaenium lineatum 550 aplopteris taeniophylla 8869 Polytaenium lineatum 875 aplopteris taeniophylla 4898 Polytaenium cajenense 279 aplopteris plurisulcata 9696 Polytaenium cajenense 906 aplopteris linearifolia 969 Polytaenium guayanense 90 aplopteris doniana 9686 Polytaenium guayanense 904 aplopteris fudzinoi 9689 Polytaenium chlorosporum 547 aplopteris dareicarpa 059 Polytaenium feei 548 aplopteris elongata 842 Polytaenium feei 258 aplopteris elongata 9075 Scoliosorus ensiformis 6020 aplopteris elongata 9070 Scoliosorus ensiformis 6652 aplopteris elongata 900 nanthacorus angustifolius 976 aplopteris elongata 909 nanthacorus angustifolius 527 aplopteris elongata 970 nanthacorus angustifolius 9704 aplopteris elongata 96 ittaria lineata 5289 aplopteris elongata 968 ittaria lineata 829 aplopteris angustifolia 8997 ittaria lineata 99 aplopteris angustifolia 9092 </ /0.80 ittaria lineata 940 aplopteris angustifolia 969 ittaria bradeorum 8559 aplopteris sikkimensis 9698 ittaria lineata 8927 aplopteris hainanensis 9690 ittaria isoetifolia 942 aplopteris ensiformis 902 ittaria isoetifolia 06 aplopteris ensiformis 94 ittaria graminifolia 9049 aplopteris ensiformis 9007 ittaria graminifolia 295 aplopteris anguste-elongata 9685 ittaria scabrida 895 aplopteris anguste-elongata 8407 ittaria scabrida 8925 aplopteris alternans 904 ittaria graminifolia 955 aplopteris alternans 904 ittaria graminifolia 554 ittaria graminifolia 88 ittaria appalachiana 94 ittaria appalachiana 9229 ersion of Record 7

7 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: R. cheesmaniae. acrocarpa (). trichoidea (2). angustissima. paradoxa (2). pumila (2) R. ruiziana (2) R. moritziana (2) R. latifolia (2) R. minima R. gardneriana () R. remota R. stipitata (). graminea (2). volkensii. humblotii. humblotii. schliebenii. guineensis (2). heterophylla. flexuosa (2). amboinensis (2). scolopendrina (). malayensis. mediosora (2). himalayensis. taeniophylla. taeniophylla. plurisulcata. linearifolia. doniana. fudzinoi. dareicarpa. elongata. elongata. elongata. elongata (2). elongata (). angustifolia (). sikkimensis. hainanensis. ensiformis. ensiformis. ensiformis. anguste-elongata. anguste-elongata. alternans (2). manniana (2). bivittata. boryana (2). obovatum. castaneum. latifolium. subfalcatum. plantagineum. parvulum (2). semicostatum (2). formosanum. henryi. coriaceum. annamense. sessilifolium (2). malgassicum. vittarioides. wallichianum (2). reticulatum. callifolium (4) P. brasilianum (2) P. citrifolium () P. lineatum () P. cajenense (2) P. guayanense (2) P. chlorosporum P. feei (2) S. ensiformis (2). angustifolius (). lineata (4). bradeorum. lineata. isoetifolia. isoetifolia. graminifolia. graminifolia. scabrida (2). graminifolia (2). graminifolia. appalachiana (2) - O M M M M I R R R R R R R M M S S S P T P P P P P S N Rheopteris aginularia ecistopteris Radiovittaria aplopteris ntrophyopsis ntrophyum Polytaenium Scoliosorus nanthacorus ittaria pinnately compound free indistinct pinnate free forked free biseriate areolate singular free pluriseriate areolate parallel free spherical slender obconic soral paraphyses absent trilete monolete - not produced I M N O P R S T leaf venation apical cells of soral paraphyses spores geographical distribution sia and the Pacific Ocean mericas frica and the Indian Ocean genus sensu Crane (997) ntrophyum aplopteris ecistopteris Monogramma nanthacorus Rheopteris Polytaenium Radiovittaria Scoliosorus netium ittaria 74 ersion of Record

8 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns resurrection of existing generic or subgeneric (in one instance) names. owever, sampling for these early studies was limited to just 2 vittarioid species and, notably, two then-recognized genera (Monogramma, Rheopteris) were not included. more recent study of the group (Ruhfel & al., 2008) revealed Monogramma to also be polyphyletic, with one element sister to Rheopteris and another rendering aplopteris paraphyletic. Nevertheless, this more recent study included only vittarioid species and also relied on a single gene (rbcl). Important parts of the family were therefore left out and, although genera (as then sampled) were typically well supported as monophyletic, some generic circumscriptions and relationships remained rather uncertain. The present study, incorporating 72 species (well over half of vittarioid diversity) and many more plastid characters, provides considerable new insight. Our phylogeny (Figs. 2, ; Electr. Suppl.: Fig. S) reveals strong support for the monophyly of ntrophyum, Radiovittaria, and ittaria, as circumscribed by Crane (997). Four additional genera in our analysis are either monotypic (nanthacorus, netium, Rheopteris) or represented by a single species (ecistopteris). owever, the four remaining genera in Crane s classification (aplopteris, Monogramma, Polytaenium, Scoliosorus) are herein resolved as being either paraphyletic or polyphyletic (Fig. ). s in Ruhfel & al. (2008), we find most of Monogramma sensu Crane (997) to be sister to Rheopteris, but we here resolve the remainder of the genus in not one, but two different places within aplopteris sensu Crane (997). Specifically, we resolve an sian / Pacific Ocean species (M. dareicarpa ook.) within a mostly sian / Pacific Ocean clade of aplopteris, whereas the frican / Indian Ocean species (M. graminea (Poir.) Schkuhr, the type of Monogramma) is sister to a purely frican / Indian Ocean aplopteris clade (Fig. ). Scoliosorus also appears in more than one place in our phylogeny. The merican species (S. ensiformis (ook.) T.Moore, the type of Scoliosorus) is strongly supported as sister to the monotypic merican genus nanthacorus, whereas three frican / Indian Ocean species previously placed in Scoliosorus form a clade sister to ntrophyum (Fig. ). Finally, we resolve Polytaenium as paraphyletic relative to the monotypic netium. Taxonomic implications. It is our opinion that supraspecific taxa ought to be monophyletic, morphologically distinguishable, and stable. lthough we understand that many existing taxa (in ferns and in other groups) fail to meet one or more of these criteria, it is our hope that by prioritizing monophyly and then diagnosability in our taxonomic revisions, we can simultaneously accomplish the goal of long-term stability (despite some requisite changes in the short-term). In this taxonomic revision of vittarioid ferns, we recognize only monophyletic genera. We also prioritize diagnosability, but we attempt to strike a balance with stability by minimizing changes relative to earlier classifications. We pay special attention to well-established monotypic genera (of which there are several in the vittarioid fern clade) and consider the merits of maintaining these taxa versus merging them with another genus. The monotypic Rheopteris, with its compound leaves and round sori (Fig. E), is unquestionably the least reduced and most distinctive vittarioid element. In this study, we find it to be sister to some of the most reduced species (previously treated in Monogramma), to which the name aginularia has been (and is) applied (see below). lthough Rheopteris shares some characteristics with this group (e.g., free venation and trilete spores), we see no compelling reason to merge the two genera. The monotypic nanthacorus is also very distinct, being the only vittarioid genus with pluriseriate areolate venation and sporangia that are restricted to submarginal commissures. Its closest relative (Scoliosorus ensiformis), although possessing the same venation and spore type, has differently shaped cells at the apices of its soral paraphyses and sporangia that are not restricted to submarginal commissures. We therefore favor the continued recognition of nanthacorus (as well as Scoliosorus; see further discussion below). netium, the third monotypic vittarioid genus following the circumscriptions of Crane (997), has traditionally been segregated based solely on the presence of sporangia between its leaf veins. But, in all other characters this taxon fits nicely within Crane s (997) definition of Polytaenium. Most notably, these two genera lack paraphyses, which are present in all other vittarioids. In our analysis, we resolve netium within Polytaenium (Fig. ); convincing support for the paraphyly of Polytaenium is not present, but taking into account morphology (and the seemingly impossible task of subdividing Polytaenium), we feel the inclusion of netium within Polytaenium is warranted. ecistopteris is not monotypic but, to date, only the most common (of three; Moran & Øllgaard, 995; Kelloff & McKee, 998) species (. pumila (Spreng.) J.Sm.) has been included in phylogenetic analyses. In our study (as in Ruhfel & al., 2008), it is well supported as sister to Radiovittaria (Fig. ). lthough these two genera both have paraphyses with obconic apical cells, this characteristic is also found in aplopteris and some species previously placed in Monogramma. dditionally, whereas Radiovittaria has biseriate areolate venation and monolete spores, ecistopteris has forked free venation and trilete spores. Thus, we maintain these genera as distinct. Fig.. ittarioid fern phylogeny, with key morphological features and geographical distributions provided for each sampled species. Phylogeny is based on maximum likelihood analysis of a four-gene (atp, chln, rbcl, rpo) plastid dataset incorporating 8 ingroup individuals (see Fig. 2). Solid branches in tree received good support (i.e., BS 70 and PP 0.95); dotted branches were poorly supported (i.e., BS < 70 or PP < 0.95). White numbers in black circles at nodes correspond to major clades / subclades discussed in the text. Samples of a given species that shared the same broad distribution were reduced to a single terminal when well supported as monophyletic (number of samples attributed to a particular terminal provided in parentheses, if more than one; see Fig. 2 for non-reduced tree and Fig. S [Electr. Suppl.] for support values). Note that species names in tree follow the generic classification proposed herein (to the right of brackets); generic assignments sensu Crane (997), indicated by letters (to the left of brackets) are provided to facilitate discussion. ersion of Record 75

9 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: Our results also support the continued recognition of ittaria as circumscribed by Crane (997). This small, mostly merican genus is well supported as monophyletic and easily recognized by the combination of biseriate areolate venation and soral paraphyses with slender apical cells. Monogramma, as circumscribed by Crane (997), and previously by Kramer (990), includes those vittarioid species with minute simple leaves having a single vein and sometimes also a few indistinct lateral veins. owever, the earliest phylogenetic studies to include members of this genus (Schuettpelz & al., 2007; Ruhfel & al., 2008) indicated that it was polyphyletic, with apparently three independent reductions in morphological complexity. Ruhfel & al. (2008) noted that one distinct element corresponded to (and included the type of) the sometimes segregated genus aginularia (Lindsay, 200). Our analysis also favors the recognition of aginularia and finds it to be strongly supported (BS = 00; PP =.00; Figs. 2, ) as sister to Rheopteris. Notably, aginularia has soral paraphyses with slender apical cells, whereas all other elements previously treated in Monogramma sensu Crane (997) have paraphyses with obconic apical cells. aginularia can also be distinguished based on its lateral veins, although these are usually indistinct (unless cleared with bleach; Ruhfel & al., 2008). The second element of Monogramma uncovered in the Ruhfel & al. (2008) analysis consisted only of M. dareicarpa, which was shown to be firmly embedded in the genus aplopteris. These authors suggested that the type of Monogramma (M. graminea) was likely also to be resolved here and pointed to the potential need to synonymize aplopteris (the younger name), based on priority, in Monogramma. owever, Lindsay & Chen (204) later argued (citing unpublished data) that M. graminea is only distantly related to M. dareicarpa; the authors thus only provided a new combination for M. dareicarpa in aplopteris. s it turns out, published data already existed to suggest that M. graminea and M. dareicarpa were not closely related. lthough the two species had not been analyzed together, their phylogenetic affinities were divergent (Schuettpelz & al., 2007; Ruhfel & al., 2008). Our analysis (Figs. 2, ) includes Monogramma dareicarpa and M. graminea simultaneously for the first time and clearly demonstrates that although both are allied to aplopteris, they occupy rather different phylogenetic positions. The sian / Pacific Ocean M. dareicarpa is closely related to the. ensiformis (Sw.) E..Crane and. elongata (Sw.) E..Crane groups within a diverse clade of predominantly sian / Pacific Ocean aplopteris species. The frican / Indian Ocean M. graminea, on the other hand, is resolved as sister to a smaller, exclusively frican / Indian Ocean clade of aplopteris (Fig. ). These two geographically divergent clades of aplopteris / Monogramma also deviate in their typical spore type, with species belonging to the former (sian / Pacific Ocean) clade generally having monolete spores (. malayensis, with trilete spores, is the exception; Fig. ), and species belonging to the latter (frican / Indian Ocean) clade having trilete spores. owever, because neither the geographical nor morphological differences are consistent among these clades, we here favor recognizing them as a single genus. The type of aplopteris (. scolopendrina (Bory) C.Presl) and that of Monogramma (M. graminea) are both resolved therein (Figs. 2, ). Monogramma is the older name (published in 809 versus 86 for aplopteris) and, based simply on the principle of priority, the correct name. That said, the vast majority of the perhaps 40 species in this group are currently treated in aplopteris; only two have ever been placed in Monogramma. For the sake of stability, we therefore propose the conservation of aplopteris over Monogramma (Chen & al., 206) and we use aplopteris as the genus name in our figures, key, and list of recognized genera herein. Of course, should our proposal fail, aplopteris would need to be replaced with Monogramma in these instances. ntrophyum was circumscribed by Crane (997) to include all Old World vittarioids with pluriseriate areolate venation and trilete spores. Old World vittarioids with pluriseriate areolate venation and monolete spores were treated in Scoliosorus. In our analysis, we find strong support (BS = 00; PP =.00; Figs. 2, ) for Crane s (997) definition of ntrophyum. owever, the situation is somewhat more complicated for those species previously placed in Scoliosorus. We find the only merican species of Scoliosorus (S. ensiformis), which is also the type, to be sister to the merican genus nanthacorus (BS = 00; PP =.00; Figs. 2, ). newly redefined Scoliosorus would therefore be monotypic; although nanthacorus is also monotypic, there is little morphological basis for combining it with Scoliosorus. The three other species of Crane s (997) Scoliosorus sampled here are all from frica and the Indian Ocean and they form a well-supported clade (BS = 00; PP =.00) that is most closely related to ntrophyum (BS = 00; PP =.00; Figs. 2, ). Because these three frican / Indian Ocean species differ from ntrophyum in having monolete (rather than trilete) spores and also have a disparate distribution (ntrophyum is essentially an sian / Pacific Ocean genus), we feel that genus-level separation is warranted. ere, we place the frican / Indian Ocean species in ntrophyopsis (Benedict) Schuettp., based on ntrophyum subg. ntrophyopsis of Benedict (907). Taxonomic summary. In all, we support the recognition of eleven vittarioid genera: nanthacorus, ntrophyopsis, ntrophyum, aplopteris, ecistopteris, Polytaenium, Radiovittaria, Rheopteris, Scoliosorus, aginularia, and ittaria. Each of these is readily diagnosable based on morphology (see key to genera and brief generic descriptions below), with micromorphological characters related to paraphyses and spores generally complementing more obvious features such as venation and the distribution of sporangia. Compared to the most recent near-comprehensive treatment (Crane, 997), the circumscriptions of five genera (nanthacorus, ntrophyum, ecistopteris, Radiovittaria, ittaria) are untouched. Rheopteris is simply brought back into consideration. netium is synonymized here within Polytaenium and the latter is thus redefined. Most species previously treated in Monogramma are placed in the genus aginularia; the remaining two (including the type of Monogramma) are incorporated into aplopteris. The name Scoliosorus is here applied only to the merican (type) species, with the frican / Indian Ocean species formerly treated in Scoliosorus now composing a new genus, ntrophyopsis 76 ersion of Record

10 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns (Benedict) Schuettp. Below, we provide a key to the currently recognized vittarioid genera, a list of genera including type information and a brief description for each genus, and five new species combinations. TXONOMY Key to currently recognized vittarioid fern genera. Leaves pinnately compound; sori round; plants of New Guinea... Rheopteris. Leaves simple, forked, or lobed; sori linear (reticulate or not); plants of frica and the Indian Ocean, the mericas, or sia and the Pacific Ocean (including New Guinea) 2 2. Leaf venation free (difficult to observe in plants with especially thick or narrow leaves unless cleared with bleach); plants generally diminutive, with leaves typically less than 0 cm long; sori usually appearing as one (continuous or discontinuous) line on each leaf (or each division if leaves forked or lobed) Leaf venation areolate (difficult to observe in plants with especially thick or narrow leaves unless cleared with bleach); plants generally larger, with leaves typically more than 0 cm long; sori always appearing as two or more lines (reticulate or not) on each leaf (or each division if leaves forked or lobed) Leaves forked or lobed; plants of the mericas ecistopteris. Leaves simple; plants of frica and the Indian Ocean or sia and the Pacific Ocean Leaf venation consisting of two or three parallel veins of equal length; apical cells of soral paraphyses spherical..... ntrophyopsis 4. Leaf venation consisting of a midrib, with or without indistinct lateral veins; apical cells of soral paraphyses slender or obconic pical cells of soral paraphyses slender; leaf venation consisting of a midrib and indistinct lateral veins; plants of sia and the Pacific Ocean... aginularia 5. pical cells of soral paraphyses obconic; leaf venation consisting of a midrib only; plants of frica and the Indian Ocean or sia and the Pacific Ocean... aplopteris 6. Leaf venation biseriate areolate, leaves with two rows of areolae (one on each side of midrib); sori restricted to submarginal commissures; leaves most often less than cm wide Leaf venation pluriseriate areolate, leaves with three or more rows of areolae; sori restricted to submarginal commissures or not; leaves most often more than cm wide pical cells of soral paraphyses slender; plants of frica and the Indian Ocean or the mericas... ittaria 7. pical cells of soral paraphyses obconic; plants of frica and the Indian Ocean, the mericas, or sia and the Pacific Ocean Rhizomes radial, leaf arrangement polystichous; plants of the mericas... Radiovittaria 8. Rhizomes dorsiventral, leaf arrangement distichous; plants of frica and the Indian Ocean or sia and the Pacific Ocean... aplopteris 9. Sori restricted to submarginal commissures; plants of the mericas... nanthacorus 9. Sori not restricted to submarginal commissures; plants of frica and the Indian Ocean, the mericas, or sia and the Pacific Ocean Soral paraphyses absent; plants of the mericas Polytaenium 0. Soral paraphyses present; plants of frica and the Indian Ocean, the mericas, or sia and the Pacific Ocean... Spores trilete; apical cells of soral paraphyses slender or spherical; plants of frica and the Indian Ocean or sia and the Pacific Ocean... ntrophyum. Spores monolete; apical cells of soral paraphyses spherical; plants of frica and the Indian Ocean or the mericas Leaves costate, midrib present; plants of the mericas..... Scoliosorus 2. Leaves ecostate, midrib absent; plants of frica and the Indian Ocean... ntrophyopsis Currently recognized vittarioid fern genera nanthacorus Underw. & Maxon in Contr. U.S. Natl. erb. 0: Type: nanthacorus angustifolius (Sw.) Underw. & Maxon ( Pteris angustifolia Sw.). simple; costate; venation pluriseriate areolate. Sori linear, restricted to submarginal commissures; soral paraphyses with slender apical cells. Spores monolete. One species in tropical merica. ntrophyopsis (Benedict) Schuettp., stat. nov. ntrophyum subg. ntrophyopsis Benedict in Bull. Torrey Bot. Club 4: Type: ntrophyopsis boryana (Willd.) Schuettp. ( emionitis boryana Willd.). simple; ecostate; venation pluriseriate areolate or, in one species, free and consisting of 2 parallel veins of equal length. Sori linear, along reticulate veins or, in one species with free venation, along the 2 parallel veins; soral paraphyses with spherical apical cells. Spores monolete. Three species in tropical frica and the Indian Ocean. ntrophyum Kaulf., Enum. Filic.: 97, Lectotype (designated by J.Sm., ist. Fil ): ntrophyum plantagineum (Cav.) Kaulf. ( emionitis plantaginea Cav.). simple; ecostate or, in a few species, partially costate; venation pluriseriate areolate. Sori linear, along reticulate veins; soral paraphyses with slender or spherical apical cells. Spores trilete. Perhaps 40 species. Most species in tropical sia and the Pacific Ocean, but at least one species in tropical frica and the Indian Ocean. ersion of Record 77

11 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: aplopteris C.Presl, Tent. Pterid.: Type: aplopteris scolopendrina (Bory) C.Presl ( Pteris scolopendrina Bory). = Pleurofossa Nakai ex.ito in J. Jap. Bot. 2: Type: Pleurofossa dareicarpa (ook.) Nakai ex.ito ( dareaecarpa ) ( Monogramma dareicarpa ook. ( dareaecarpa )). simple or, in a few species, forked; costate; venation biseriate areolate or, in two species, consisting of a single vein. Sori linear, restricted to submarginal commissures or, in two species, along the single vein; soral paraphyses with obconic apical cells. Spores monolete or trilete. Perhaps 40 species. Most species in tropical sia and the Pacific Ocean, but about ten species in tropical frica and the Indian Ocean. For the sake of stability, we propose the conservation of aplopteris over Monogramma Comm. ex Schkuhr (24 Kl. Linn. Pfl.-Syst.: ), which, based on the principle of priority, would be the correct name for this group (Chen & al., 206). We use aplopteris as the genus name here, as well as in our figures and key; should our proposal fail, aplopteris would need to be replaced with Monogramma in these instances. ecistopteris J.Sm. in London J. Bot. : Type: ecistopteris pumila (Spreng.) J.Sm. ( Gymnogramma pumila Spreng.). forked or, in one species, lobed; ecostate; venation free, veins forked. Sori linear, along vein ends; soral paraphyses with obconic apical cells. Spores trilete. Three species in tropical merica. Polytaenium Desv. in Mém. Soc. Linn. Paris 6: 74, Type: Polytaenium lanceolatum (Sw.) Desv. ( ittaria lanceolata Sw.). = netium Splitg. in Tijdschr. Natuurl. Gesch. Physiol. 7: Type: netium citrifolium (L.) Splitg. ( crostichum citrifolium L.). simple; costate; venation pluriseriate areolate. Sori linear, along reticulate veins and, in one species, sporangia also scattered between veins; soral paraphyses absent. Spores trilete. bout ten species in tropical merica. Radiovittaria (Benedict) E..Crane in Syst. Bot. 22: ( 997 ) ittaria subg. Radiovittaria Benedict in Bull. Torrey Bot. Club 8: Type: Radiovittaria remota (Fée) E..Crane ( ittaria remota Fée). Rhizome radial; leaf arrangement polystichous. Leaves simple; costate; venation biseriate areolate. Sori linear, restricted to submarginal commissures; soral paraphyses with obconic apical cells. Spores monolete. bout ten species in tropical merica. Rheopteris lston in Nova Guinea, n.s., 7: Type: Rheopteris cheesmaniae lston. pinnately compound; pinnae ecostate; venation free, veins forked. Sori round, at vein ends; soral paraphyses with spherical apical cells. Spores trilete. One species in New Guinea. Scoliosorus T.Moore, Index Fil.: xxix. 857 Type: Scoliosorus ensiformis (ook.) T.Moore ( ntrophyum ensiforme ook.). simple; costate; venation pluriseriate areolate. Sori linear, along reticulate veins; soral paraphyses with spherical apical cells. Spores monolete. One species in tropical merica. aginularia Fée, Mém. Foug. : Type: aginularia trichoidea Fée. = Diclidopteris Brack., U.S. Expl. Exped., Filic. [= in Wilkes, U.S. Expl. Exped. 6]: Type: Diclidopteris angustissima Brack. simple; costate; venation free, consisting of a midrib and indistinct lateral veins. Sori linear, along lateral veins, usually appearing as one (continuous or discontinuous) line; soral paraphyses with slender apical cells. Spores trilete. Four species in tropical sia and the Pacific Ocean. ittaria Sm. in Mém. cad. Roy. Sci. (Turin) 5: 4, t. 9, fig Type: ittaria lineata (L.) Sm. ( Pteris lineata L.). simple; costate; venation biseriate areolate. Sori linear, restricted to submarginal commissures; soral paraphyses with slender apical cells. Spores monolete or trilete. Seven species. Most species in tropical merica, but one in tropical frica and the Indian Ocean and one restricted to temperate North merica. New combinations ntrophyopsis bivittata (C.Chr.) Schuettp., comb. nov. ntrophyum bivittatum C.Chr. in Bonaparte, Notes Ptéridol. 6: 0, pl ntrophyopsis boryana (Willd.) Schuettp., comb. nov. emionitis boryana Willd., Sp. Pl. 5(): ntrophyum boryanum (Willd.) Spreng., Syst. eg. 4(): Scoliosorus boryanus (Willd.) E..Crane in Syst. Bot. 22: ( 998 ) ( boryanum ). ntrophyopsis manniana (ook.) Schuettp., comb. nov. ntrophyum mannianum ook., Sec. Cent. Ferns: t Scoliosorus mannianus (ook.) E..Crane in Syst. Bot. 22: ( 997 ) ( mannianum ). aplopteris schliebenii (Reimers) Schuettp., comb. nov. ittaria schliebenii Reimers in Notizbl. Bot. Gart. Berlin- Dahlem : ersion of Record

12 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns Polytaenium citrifolium (L.) Schuettp., comb. nov. crostichum citrifolium L., Sp. Pl.: netium citrifolium (L.) Splitg. in Tijdschr. Natuurl. Gesch. Physiol 7: ntrophyum citrifolium (L.) Fée, Mém. Foug. 4: emionitis citrifolia (L.) ook., Sp. Fil. 5: Pteridanetium citrifolium (L.) Copel., Gen. Fil.: CKNOWLEDGEMENTS We are grateful to Sally Chambers, Maarten Christenhusz, Thomas Janssen, Fay-Wei Li, Fernando Matos, Joel Nitta, Jefferson Prado, Tom Ranker, Carl Rothfels, Germinal Rouhan, lan Smith, Michael Sundue, and George Yatskievych for their assistance with the acquisition of some material for this study. We also thank Stuart Lindsay for drawing our attention to the exceptional spores of aplopteris malayensis and Jefferson Prado for supplying nomenclatural expertise. Three anonymous reviewers and the editors provided helpful comments. Fay-Wei Li, Germinal Rouhan, Michael Sundue (www. fernsoftheworld.com), and Wayne Takeuchi kindly provided images. Portions of the laboratory work were conducted in and with the support of the L..B. facilities of the National Museum of Natural istory. Funding for this research was provided in part by the United States National Science Foundation (award DEB-4058 to E.S. and award DEB-4564 to K.M.P. and L..). LITERTURE CITED Benedict, R.C The genus ntrophyum I. Synopsis of subgenera, and the merican species. Bull. Torrey Bot. Club 4: Benedict, R.C. 9. The genera of the fern tribe ittarieae: Their external morphology, venation and relationships. Bull. Torrey Bot. Club 8: Beck, J.B., lexander, P.J., llphin, L., l-shehbaz, I.., Rushworth, C., Bailey, C.D. & Windham, M.D. 20. Does hybridization drive the transition to asexuality in diploid Boechera? 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Taxon 65: Ching, R.C On natural classification of the family Polypodiaceae. Sunyatsenia 5: Christenhusz, M.J.M. & Chase, M.W Trends and concepts in fern classification. nn. Bot. (Oxford) : Cochran,., Prado, J. & Schuettpelz, E Tryonia, a new taenitidoid fern genus segregated from Jamesonia and Eriosorus (Pteridaceae). PhytoKeys 5: Copeland, E.B Genera Filicum. Waltham: Chronica Botanica. Crane, E revised circumscription of the genera of the fern family ittariaceae. Syst. Bot. 22: Crane, E.., Farrar, D.R. & Wendel, J.F Phylogeny of the ittariaceae: Convergent simplification leads to a polyphyletic ittaria. mer. Fern J. 85: Doyle, J.J. & Doyle, J.L rapid DN isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull. Bot. Soc. mer. 9: 5. Edgar, R.C MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucl. cids Res. 2: asebe, M., Wolf, P.G., Pryer, K.M., Ueda, K., Ito, M., Sano, R., Gastony, G.J., Yokoyama, J., Manhart, J.R., Murakami, N., Crane, E.., aufler, C.. & auk, W.D Fern phylogeny based on rbcl nucleotide sequences. mer. Fern J. 85: uelsenbeck, J.P. & Ronquist, F MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 7: Kelloff, C. & McKee, G new species of ecistopteris from Guyana, South merica. mer. Fern J. 88: Kramer, K.U ittariaceae. Pp in: Kramer, K.U. & Green P.S. (eds.), The families and genera of vascular plants, vol., Pteridophytes and gymnosperms. Berlin: Springer. Larsson, liiew: fast and lightweight alignment viewer and editor for large data sets. Bioinformatics 0: Lindsay, S Considerations for a revision of the fern family ittariaceae for Flora Malesiana. Telopea 0: Lindsay, S. & Chen, C.W Three new combinations in aplopteris (Pteridaceae subfam. ittarioideae). Gard. Bull. Singapore 66: Lu, J.-M., Wen, J., Lutz, S., Wang, Y.-P. & Li, D.-Z. 20. Phylogenetic relationships of Chinese diantum based on five plastid markers. J. Pl. Res. 25: Moran, R.C. & Øllgaard, B Six new species of ferns (Polypodiopsida) from Ecuador. Nordic J. Bot. 5: Prado, J., Rodrigues, C.D.N., Salatino,. & Salatino, M.L.F Phylogenetic relationships among Pteridaceae, including Brazilian species, inferred from rbcl sequences. Taxon 56: Pryer, K.M., uiet, L., Li, F.-W., Rothfels, C.J. & Schuettpelz, E Maidenhair ferns diantum are indeed monophyletic and sister to shoestring ferns vittarioids (Pteridaceae). Syst. Bot. 4: Rambaut,., Suchard, M.., Xie, D. & Drummond,.J Tracer, version.6. Ronquist, F. & uelsenbeck, J.P MrBayes : Bayesian phylogenetic inference under mixed models. Bioinformatics 9: Rothfels, C.J. & Schuettpelz, E ccelerated rate of molecular evolution for vittarioid ferns is strong and not driven by selection. Syst. Biol. 6: ersion of Record 79

13 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: Rothfels, C.J., Sundue, M.., Kuo, L.-Y., Larsson,., Kato, M., Schuettpelz, E. & Pryer, K.M revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 6: Rothfels, C.J., Li, F.-W., Sigel, E.M., uiet, L., Larsson,., Burge, D.O., Ruhsam, M., Deyholos, M., Soltis, D.E., Stewart, C.N., Jr., Shaw, S.W., Pokorny, L., Chen, T., depamphilis, C., DeGironimo, L., Chen, L., Wei, X., Sun, X., Korall, P., Stevenson, D.W., Graham, S.W., Wong, G.K.-S. & Pryer, K.M The evolutionary history of ferns inferred from 25 low-copy nuclear genes. mer. J. Bot. 02: Ruhfel, B., Lindsay, S. & Davis, C.C Phylogenetic placement of Rheopteris and the polyphyly of Monogramma (Pteridaceae s.l.): Evidence from rbcl sequence data. Syst. Bot. : Schneider,., Smith,.R. & Pryer, K.M Is morphology really at odds with molecules in estimating fern phylogeny? Syst. Bot. 4: Schuettpelz, E. & Pryer, K.M Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: Schuettpelz, E., Korall, P. & Pryer, K.M Plastid atp data provide improved support for deep relationships among ferns. Taxon 55: Schuettpelz, E., Schneider,., uiet, L., Windham, M.D. & Pryer, K.M molecular phylogeny of the fern family Pteridaceae: ssessing overall relationships and the affinities of previously unsampled genera. Molec. Phylogen. Evol. 44: Schuettpelz, E., Grusz,.L., Windham, M.D. & Pryer, K.M The utility of nuclear gapcp in resolving polyploid fern origins. Syst. Bot. : Smith,.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider,. & Wolf, P.G classification for extant ferns. Taxon 55: Stamatakis, RxML version 8: tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 0: 2. Sundue, M Silica bodies and their systematic implications in Pteridaceae (Pteridophyta). Bot. J. Linn. Soc. 6: Sundue, M.. & Rothfels, C.J Stasis and convergence characterize morphological evolution in eupolypod II ferns. nn. Bot. (Oxford) : Tryon, R.M. & Tryon,.F Ferns and allied plants with special reference to tropical merica. Berlin: Springer. Willams, S critical examination of the ittarieae with a view to their systematic comparison. Trans. Roy. Soc. Edinburgh 55: ppendix. Individuals sampled in this study of vittarioid ferns. For each individual, the species name, Fern Lab Database ( edu) voucher number, voucher information, place of origin, and GenBank accession numbers (atp, chln, rbcl, rpo) are provided. n n-dash ( ) indicates unavailable information; new sequences are prefixed with KX. diantum andicola Liebm., 5549, C.J. Rothfels 264 (DUKE), Costa Rica (San José), KU4724, KU4725, KU47272, KU47280; diantum capillusveneris L., 4609, L. uiet 04 (UC), United States (California), KU47244, KU47252, KU4727, KU4728; diantum formosum R.Br., 4602,.R. Smith s.n. (UC), Cultivated ( ), KC98447, KU47257, KC984520, KU47289; diantum malesianum J.Ghatak, 2506, L. uiet (UC), Cultivated ( ), KX6557, KU47258, EF4522, KU4729; diantum peruvianum Klotzsch, 2507, L. uiet 0 (UC), Cultivated ( ), KX6558, KU47259, EF452, KU47292; diantum tenerum Sw., 2504, L. uiet 07 (UC), Cultivated ( ), KX6559, KU47260, EF4524, KU47294; diantum tetraphyllum umb. & Bonpl. ex Willd., 2505, L. uiet 05 (UC), Cultivated ( ), EF45207, KU4726, EF4525, KU47295; nanthacorus angustifolius (Sw.) Underw. & Maxon, 527, F. Matos (DUKE), Costa Rica (eredia), KU57449, KU57485, KU57577, KU576; nanthacorus angustifolius (Sw.) Underw. & Maxon, 976,. Garrido 2 (MO), Nicaragua (Río San Juan), KX6560,, KX64950, ; nanthacorus angustifolius (Sw.) Underw. & Maxon, 9704, P.M. Peterson 684 (US), Panama (Bocas del Toro), KX656, KX6506, KX6495, KX6485; ntrophyopsis bivittata (C.Chr.) Schuettp., 996, J. Rabenantoandro 52 (MO), Madagascar (ntsiranana), KX6562,, KX64952, ; ntrophyopsis boryana (Willd.) Schuettp., 9076, J. Kluge 8047 (UC), Madagascar (Sava), KX65246, KX65064, KX6495, KX64854; ntrophyopsis boryana (Willd.) Schuettp., 970, J.S. Miller 54 (US), Madagascar (ntsiranana), KX656, KX65065, KX64954, KX64855; ntrophyopsis manniana (ook.) Schuettp., 940, L. Festo 2008 (MO), Tanzania (Iringa), KX6564, KX65066, KX64955, KX64856; ntrophyopsis manniana (ook.) Schuettp., 9702, I. Friis 98 (US), Ethiopia ( ), KX6565, KX65067, KX64956, KX64857; ntrophyum annamense Tardieu & C.Chr., 99, S.K. Wu 96 (MO), Laos (Kouankhanh), KX6572, KX6507, KX6496, KX6486; ntrophyum annamense Tardieu & C.Chr., 967, W.. Wu 956 (TIF), China (ainan), KX6566, KX65068, KX64957, KX64858; ntrophyum callifolium Blume, 408, E. Schuettpelz 668 (DUKE), Malaysia (Selangor), KX6567, KX65069, KX64958, KX64859; ntrophyum callifolium Blume, 908, D. Karger 875 (UC), Indonesia (Sulawesi), KX6568, KX65070, KX64959, KX64860; ntrophyum callifolium Blume, 99, W. Sugong 826 (MO), Laos (Saraven), KX6569,, KX64960, ; ntrophyum callifolium Blume, 9674, C.W. Chen SITW0065 (TIF), Solomon Islands (Guadalcanal), KX6570, KX6507, KX6496, KX6486; ntrophyum castaneum.ito, 9675, C.W. Chen 75 (TIF), Taiwan (sinchu), KX657, KX65072, KX64962, KX64862; ntrophyum formosanum ieron., 8409, T. Ranker 2068 (COLO), Taiwan (Taipei), KU5745, KU57487, KU57579, KU5765; ntrophyum henryi ieron., 9677, L.Y. Kuo 4 (TIF), China (Yunnan), KX657, KX65074, KX64964, KX64864; ntrophyum latifolium Blume, 078, T. Ranker 774 (COLO), Papua New Guinea ( ), KX6574, KU4726, EF4528, KU47297; ntrophyum malgassicum C.Chr., 998, F. Rakotondrainibe 70 (MO), Madagascar (ntsiranana), KX6575,, KX64965, ; ntrophyum obovatum Baker, 842, T. Ranker 2082 (COLO), Taiwan (Nantou), KX6576, KX65075, KX64966, KX64865; ntrophyum parvulum Blume, 900, D. Karger (UC), Indonesia (Lombok), KX65247, KX65076, KX64967, KX64866; ntrophyum parvulum Blume, 9679, C.W. Chen 222 (TIF), Taiwan (Chiayi), KX6577, KX65077, KX64968, KX64867; ntrophyum plantagineum (Cav.) Kaulf., 9680, C.W. Chen SITW028 (TIF), Solomon Islands (Western), KX6578, KX65078, KX64969, KX64868; ntrophyum reticulatum (G.Forst.) Kaulf., 844, T. Ranker 948 (COLO), French Polynesia (Moorea), KX6579, KX65079, KX64970, KX64869; ntrophyum semicostatum Blume, 845, T. Ranker 262 (SN), Malaysia (Sabah), KX6580, KX65080, KX6497, KX64870; ntrophyum semicostatum Blume, 9066, J. Kluge 75 (UC), Indonesia (Sulawesi), KX65248, KX6508, KX64972, KX6487; ntrophyum sessilifolium (Cav.) Spreng., 8999, D. Cicuzza 927 (UC), Indonesia (Sulawesi), KX65249, KX65082, KX6497, KX64872; ntrophyum sessilifolium (Cav.) Spreng., 968, C.W. Chen 502 (TIF), Taiwan (Taitung), KX658, KX6508, KX64974, KX6487; ntrophyum subfalcatum Brack., 9682, C.W. Chen SITW0 (TIF), Solomon Islands (Western), KX6582, KX65084, KX64975, KX64874; ntrophyum vittarioides Baker, 968, Lao (TIF), China (Guangxi), KX658, KX65085, KX64976, KX64875; ntrophyum wallichianum M.G.Gilbert & X.C.Zhang, 995, W. Sugong 2585 (MO), Laos (Louangphrabang), KX6584, KX65086, KX64977, KX64876; ntrophyum wallichianum M.G.Gilbert & X.C.Zhang, 9676, C.R. Fraser-Jenkins FN2 (TIF), Nepal ( ), KX6585, KX65087, KX64978, KX64877; aplopteris alternans (Copel.) S.Linds. & C.W.Chen, 904, D. Karger 685 (UC), Philippines ( ), KX6586, KX65088, KX64979, KX64878; aplopteris alternans (Copel.) S.Linds. & C.W.Chen, 904, D. Karger 508 (UC), Philippines (Bukiknon), KX6556, KX65089, KX64980, KX64879; aplopteris amboinensis (Fée) X.C.Zhang, 97, Wuzhishan Fern Survey 22 (MO), China (ainan), KX6587, KX65090, KX6498, ; aplopteris amboinensis (Fée) X.C.Zhang, 9684, C.W. Chen 2588 (TIF), ietnam (Lam Dong), KX6588, KX6509, KX64982, KX64880; aplopteris anguste-elongata (ayata) E..Crane, 8407, T. Ranker 202 (COLO), Taiwan (Ilan), KX65250, KX65092, KX6498, KX6488; aplopteris anguste-elongata (ayata) E..Crane, 9685, 720 ersion of Record

14 TXON 65 (4) ugust 206: Schuettpelz & al. Revised classification of vittarioid ferns ppendix. Continued. C.W. Chen 482 (TIF), Taiwan (Taipei), KX6589, KX6509, KX64984, KX64882; aplopteris angustifolia (Blume) E..Crane, 8997, D. Cicuzza 88 (UC), Indonesia (Sulawesi), KX6525, KX65094, KX64985, KX6488; aplopteris angustifolia (Blume) E..Crane, 9092, M. Kessler 6 (UC), Malaysia ( ), KX6590, KX65095, KX64986, KX64884; aplopteris angustifolia (Blume) E..Crane, 969, Wuzhishan Fern Survey 26 (MO), China (ainan), KX659, KX65096, KX64987, KX64885; aplopteris dareicarpa (ook.) S.Linds. & C.W.Chen, 059, P.J. Darbyshire 802 (US), Papua New Guinea ( ), KX6592, KX65097, KX64988, KX64886; aplopteris doniana (Mett. ex ieron.) E..Crane, 9686, C.W. Chen 2680 (TIF), ietnam (Lam Dong), KX659, KX65098, KX64989, KX64887; aplopteris elongata (Sw.) E..Crane, 842, T. Ranker 9 (COLO), French Polynesia (Moorea), KX6594, KX65099, KX64990, KX64888; aplopteris elongata (Sw.) E..Crane, 900, D. Cicuzza 977 (Z), Indonesia (Sulawesi), KX65252, KX6500, KX6499, KX64889; aplopteris elongata (Sw.) E..Crane, 9070, J. Kluge 7677 (UC), Comoros (Grand Comore), KX6595, KX650, KX64992, KX64890; aplopteris elongata (Sw.) E..Crane, 9075, J. Kluge 7865 (UC), Madagascar ( ), KX6525, KX6502, KX6499, KX6489; aplopteris elongata (Sw.) E..Crane, 909, M. Kessler 62 (UC), Malaysia ( ), KX65254, KX650, KX64994, KX64892; aplopteris elongata (Sw.) E..Crane, 96, S.K. Wu 94 (MO), Laos (ientiane), KX6596, KX6504, KX64995, ; aplopteris elongata (Sw.) E..Crane, 968, Wuzhishan Fern Survey 26 (MO), China (ainan),,, KX64996, ; aplopteris elongata (Sw.) E..Crane, 970, S.K. Wu 864 (MO), ietnam (a Giang), KX6597, KX6505, KX64997, KX6489; aplopteris ensiformis (Sw.) E..Crane, 9007, D. Karger (UC), Philippines (Bukiknon), KX6598, KX6506, KX64998, ; aplopteris ensiformis (Sw.) E..Crane, 902, D. Karger 40 (UC), Indonesia (Seram),,, KX64999, ; aplopteris ensiformis (Sw.) E..Crane, 94,. van der Werff 2768 (MO), Madagascar (Toamasina), KX6599, KX6507, KX65000, ; aplopteris flexuosa (Fée) E..Crane, 4786, E. Schuettpelz 04 (DUKE), Taiwan (Nantou), KX65255, KX6508, KX6500, KX64894; aplopteris flexuosa (Fée) E..Crane, 9688, L.Y. Kuo 265 (TIF), China (Yunnan), KX65200, KX6509, KX65002, KX64895; aplopteris fudzinoi (Makino) E..Crane, 9689, L.Y. Kuo 2225 (TIF), China (Sichuan), KX6520, KX650, KX6500, KX64896; aplopteris graminea (Poir.) comb. ined. (= Monogramma graminea (Poir.) Schkuhr), 548, T. Janssen 2692 (P), France (Reunion), KX65202, KU47268, EF45257, KU4704; aplopteris graminea (Poir.) comb. ined. (= Monogramma graminea (Poir.) Schkuhr), 058, G. Rouhan 4 (P), Madagascar ( ),,, KX65004, ; aplopteris guineensis (Desv.) E..Crane, 959, W. Kindeketa 850 (MO), Tanzania (Kilimanjaro), KX6520, KX65, KX65005, KX64897; aplopteris guineensis (Desv.) E..Crane, 960, R. Pérez iso 72 (MO), Equatorial Guinea (Wele-Nzas), KX65204, KX652, KX65006, KX64898; aplopteris hainanensis (C.Christensen ex Ching) E..Crane, 9690, W.. Wu 959 (TIF), China (ainan), KX65205, KX65, KX65007, KX64899; aplopteris heterophylla C.W.Chen, Y..Chang & Y.C.Liu, 969, W.. Wu 08 (TIF), China (ainan), KX65206, KX654, KX65008, KX64900; aplopteris himalayensis (Ching) E..Crane, 9692, C.R. Fraser-Jenkins 0640 (TIF), Nepal ( ),,, KX65009, KX6490; aplopteris humblotii (ieron.) S.Linds. & C.W.Chen, 945, P. ntilahimena 77 (MO), Madagascar (Toamasina), KX65207, KX655, KX6500, KX64902; aplopteris humblotii (ieron.) S.Linds. & C.W.Chen, 946, F. Rakotondrainibe 554 (MO), Madagascar (ntsiranana), KX65208,, KX650, ; aplopteris linearifolia (Ching) X.C.Zhang, 969, Y.C. Liu 9457 (TIF), China (Sichuan), KX65209, KX656, KX6502, KX6490; aplopteris malayensis (olttum) E..Crane, 44, E. Schuettpelz 728 (DUKE), Malaysia (Pahang), KX65256, KX657, KX650, KX64904; aplopteris mediosora (ayata) X.C.Zhang, 9694, C.R. Fraser-Jenkins FN (TIF), Nepal ( ), KX6520, KX658, KX6504, KX64905; aplopteris mediosora (ayata) X.C.Zhang, 9695, C.W. Chen 492 (TIF), Taiwan (Nantou), KX652, KX659, KX6505, KX64906; aplopteris plurisulcata (Ching) X.C.Zhang, 9696, Z.R. e 2000 (TIF), China (Yunnan), KX6522, KX6520, KX6506, KX64907; aplopteris schliebenii (Reimers) Schuettp., 9407, W. R.Q. Luke 55 (MO), Tanzania (Iringa), KX652, KX652, KX6507, KX64908; aplopteris scolopendrina (Bory) C.Presl, 905, D. Karger 50 (UC), Philippines (Bukiknon),,, KX6508, ; aplopteris scolopendrina (Bory) C.Presl, 906, J. Kluge 7052 (GOET), Indonesia (Sulawesi), KX65257, KX6522, KX6509, KX64909; aplopteris scolopendrina (Bory) C.Presl, 9697, C.W. Chen SITW047 (TIF), Solomon Islands (Western), KX6524,,, KX6490; aplopteris sikkimensis (Kuhn) E..Crane, 9698, C.R. Fraser-Jenkins 0779 (TIF), Nepal ( ), KX6525,, KX65020, ; aplopteris taeniophylla (Copel.) E..Crane, 4898, E. Schuettpelz 55 (DUKE), Taiwan (Taichung), KX65258, KX652, KX6502, KX649; aplopteris taeniophylla (Copel.) E..Crane, 8869, F.W. Li 440 (DUKE), Taiwan (Nantou), KX6526, KX6524, KX65022, KX6492; aplopteris volkensii (ieron.) E..Crane, 94, L. Festo 97 (MO), Tanzania (Morogoro), KX6527, KX6525, KX6502, ; ecistopteris pumila (Spreng.) J.Sm., 278, M. J.M. Christenhusz 976 (TUR), Guadeloupe (Sofaia), EF452097, KU47266, KC984524, KU470; ecistopteris pumila (Spreng.) J.Sm., 55, F. Matos (DUKE), Costa Rica (eredia), KX6528, KX6526, KX65024, KX649; Polytaenium brasilianum (Desv.) Benedict, 9087, M. Kessler 2940 (UC), Bolivia (Cochabamba), KX6529, KX6527, KX65025, KX6494; Polytaenium brasilianum (Desv.) Benedict, 9090, M. Kessler 5 (UC), Bolivia (Cochabamba), KX65220, KX6528, KX65026, KX6495; Polytaenium cajenense (Desv.) Benedict, 279, E. Schuettpelz 2 (DUKE), Ecuador (Napo), KX65259, KX6529, KX65027, KX6496; Polytaenium cajenense (Desv.) Benedict, 906, P. Weigelt 9065 (LPB), Bolivia (La Paz), KX6522, KX650, KX65028, KX6497; Polytaenium chlorosporum (Mickel & Beitel) E..Crane, 547, C.J. Rothfels (DUKE), Costa Rica (Puntarenas), KX65260, KX65, KX65029, KX6498; Polytaenium citrifolium (L.) Schuettp., 9, M. J.M. Christenhusz 4076 (TUR), Guadeloupe ( ), EF452075, KU47262, KC98452, KU47296; Polytaenium citrifolium (L.) Schuettp., 5, C.J. Rothfels (DUKE), Costa Rica (eredia), KX65222, KX652, KX6500, KX6499; Polytaenium citrifolium (L.) Schuettp., 906, I. Jiménez 2007 (UC), Bolivia (Pando), KU57450,, KU57578, KU5764; Polytaenium feei (W.Schaffn. ex Fée) Maxon, 258, M. J.M. Christenhusz 95 (TUR), Guadeloupe (Sofaia), KX6526,, KX650, KX64920; Polytaenium feei (W.Schaffn. ex Fée) Maxon, 548, C.J. Rothfels (DUKE), Costa Rica (eredia), KX6522,, KX6502, KX6492; Polytaenium guayanense (ieron.) lston, 90, P. Weigelt 9007 (LPB), Bolivia (Cochabamba), KX65224, KX65, KX650, KX64922; Polytaenium guayanense (ieron.) lston, 904, P. Weigelt 904 (LPB), Bolivia (Cochabamba), KX65225, KX654, KX6504, KX6492; Polytaenium lineatum (Sw.) J.Sm., 550, K.M. Pryer (DUKE), Costa Rica (Puntarenas), KX65226, KX655, KX6505, KX64924; Polytaenium lineatum (Sw.) J.Sm., 875, E. Schuettpelz 440 (SP), Brazil (Rio de Janeiro), KU57452, KU57488, KU57580, KU5766; Polytaenium lineatum (Sw.) J.Sm., 9062, I. Jiménez 29 (UC), Bolivia (Tarija), KX65227, KX656, KX6506, KX64925; Radiovittaria gardneriana (Fée) E..Crane, 445, E. Schuettpelz 929 (DUKE), Ecuador (Napo), KX65228, KX657, KX6507, KX64926; Radiovittaria gardneriana (Fée) E..Crane, 870, E. Schuettpelz 46 (SP), Brazil (Rio de Janeiro),, KX658, KX6508, KX64927; Radiovittaria gardneriana (Fée) E..Crane, 905, I. Jiménez 90 (LPB), Bolivia (La Paz), KX65229, KX659, KX6509, KX64928; Radiovittaria latifolia (Benedict) E..Crane, 9052, I. Jiménez 069 (UC), Bolivia (La Paz), KX6520, KX6540, KX65040, KX64929; Radiovittaria latifolia (Benedict) E..Crane, 9058, I. Jiménez 525 (UC), Bolivia (Cochabamba), KX652, KX654, KX6504, KX6490; Radiovittaria minima (Baker) E..Crane, 855, J. Nitta 806 (UC), Costa Rica (lajuela), KX6522, KX6542, KX65042, KX649; Radiovittaria moritziana (Mett.) E..Crane, 9046, I. Jiménez 689 (UC), Bolivia (La Paz), KX652, KX654, KX6504, KX6492; Radiovittaria moritziana (Mett.) E..Crane, 9059, I. Jiménez 606 (UC), Bolivia (Cochabamba), KX6524, KX6544, KX65044, KX649; Radiovittaria remota (Fée) E..Crane, 854, J. Nitta 72 (UC), Costa Rica (lajuela),, KX6545,, KX6494; Radiovittaria ruiziana (Fée) E..Crane, 977, J.P. ltamirano 5 (MO), Bolivia (Cochabamba), KX6525, KX6546, KX65045, KX6495; Radiovittaria ruiziana (Fée) E..Crane, 979, R. Núnez 8 (MO), Bolivia (Santa Cruz), KX6526, KX6547, KX65046, KX6496; Radiovittaria stipitata (Kunze) E..Crane, 249, E. Schuettpelz 25 (DUKE), Ecuador (Zamora-Chinchipe), KX6527, KX6548, KX65047, KX6497; Radiovittaria stipitata (Kunze) E..Crane, 55, C.J. Rothfels (DUKE), Costa Rica (eredia), KX6528, KX6549, KX65048, KX6498; Radiovittaria stipitata (Kunze) E..Crane, 9045,. uaylla 400 (UC), Bolivia (Cochabamba), KX6529, KX6550, KX65049, KX6499; Rheopteris cheesmaniae lston, 7, J.R. Croft 749 (), Papua New Guinea ( ), EF45226, KU47270, EF45276, KU470; Scoliosorus ensiformis (ook.) T.Moore, 6020, M. Sundue 676 (DUKE), Costa Rica (lajuela), KU5745, KU57489, KU5758, KU5767; Scoliosorus ensiformis (ook.) T.Moore, 6652, C.J. Rothfels 254 (DUKE), Mexico (Guerrero), KX65240, KX655, KX65050, KX64940; aginularia acrocarpa olttum, 75, T. Ranker 778 (COLO), Papua New Guinea ( ), KC98445,, EF45256, ; aginularia acrocarpa olttum, 9005, D. Karger 220 (UC), Philippines ( ), KX65262,, KX6505, ; aginularia acrocarpa olttum, 900, M. Kessler 4254 (Z), ustralia (Queensland), KX6526,, KX65052, ; aginularia angustissima (Brack.) Mett., 9705,.C. Smith 5806 (US), Fiji ( ), KX6524,, KX6505, KX6494; aginularia paradoxa (Fée) Mett. ex Miq., 906, D. Karger 548 (UC), Philippines (Bukiknon), KX65264, ersion of Record 72

15 Schuettpelz & al. Revised classification of vittarioid ferns TXON 65 (4) ugust 206: ppendix. Continued., KX65054, KX64942; aginularia paradoxa (Fée) Mett. ex Miq., 9699, C.W. Chen 64 (TIF), Taiwan (Taitung), KX65242,, KX65055, KX6494; aginularia trichoidea Fée, 4668, F.C. ou 74 (UC), China (ainan), KX6524,, KX65056, KX64944; aginularia trichoidea Fée, 909, M. Kessler 60 (UC), Malaysia ( ), KX65265,, KX65057, KX64945; ittaria appalachiana Farrar & Mickel, 9229, E. Schuettpelz 482 (WNC), United States (North Carolina), KU57454, KU57490, KU57582, KU5768; ittaria appalachiana Farrar & Mickel, 94, S. Stevens NY-p5s2 (PUL), United States (New York), KU57460, KU57495, KU57588, KU57624; ittaria bradeorum Rosenst., 8559, J. Nitta 84 (UC), Costa Rica (lajuela), KU57472, KU57507, KU57600, KU5766; ittaria graminifolia Kaulf., 295, E. Schuettpelz 227 (DUKE), Ecuador (Zamora-Chinchipe), EF45228, KU4727, KU47279, KU4702; ittaria graminifolia Kaulf., 554, C.J. Rothfels (DUKE), Costa Rica (Puntarenas), KX65266, KX6552, KX65058, KX64946; ittaria graminifolia Kaulf., 88, E. Schuettpelz 446 (SP), Brazil (Rio de Janeiro), KU57475, KU5750, KU5760, KU5769; ittaria graminifolia Kaulf., 9049, I. Jiménez 867 (LPB), Bolivia (La Paz), KX65244, KX655, KX65059, KX64947; ittaria graminifolia Kaulf., 955,. Reyes-García 720 (MO), Mexico (Chiapas), KX65245, KX6554, KX65060, KX64948; ittaria isoetifolia Bory, 942, F. Rakotondrainibe 202 (MO), Madagascar (Fort Dauphin),, KU575, KU57606, KU57642; ittaria isoetifolia Bory, 06, T. Janssen 2920 (MO), Madagascar ( ), KU57478, KU5754,, KU5764; ittaria lineata (L.) Sm., 5289, C.J. Rothfels (DUKE), Costa Rica (eredia), KU57479, KU5755, KU57607, KU57644; ittaria lineata (L.) Sm., 829, C.J. Rothfels 4008 (DUKE), United States (Florida), KU57480, KU5756, KU57608, KU57645; ittaria lineata (L.) Sm., 8927, J. Prado 2248 (SP), Brazil (Sao Paulo), KX65267, KX6555, KX6506, KX64949; ittaria lineata (L.) Sm., 99, D.E. tha 0 (MO), Belize (Cayo),,, KX65062, ; ittaria lineata (L.) Sm., 940, G. McPherson 2097 (MO), Panama ( ), KU57482, KU5758, KU5760, KU57647; ittaria scabrida Klotzsch ex Fée, 895, J. Prado 2249 (SP), Brazil (Sao Paulo), KU5748, KU5759, KU576, KU57648; ittaria scabrida Klotzsch ex Fée, 8925, J. Prado 226b (SP), Brazil (Sao Paulo), KU57484, KU57520, KU5762, KU ersion of Record

16 ol. 65 (4) ugust 206 International Journal of Taxonomy, Phylogeny and Evolution Electronic Supplement to revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data Eric Schuettpelz, Cheng-Wei Chen, Michael Kessler, Jerald B. Pinson, Gabriel Johnson, lex Davila, lyssa T. Cochran, Layne uiet & Kathleen M. Pryer Taxon 65: