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INTRODUCTION

Hasebe & al. (1994, 1995) and Pryer & al. (1995) first investigated fern phylogeny based on rbcL, and morphology plus rbcL, respectively. Hasebe & al. (1994) showed that Poly-podiaceae Bercht. & J. Presl plus Grammitidaceae Newman form a monophyletic group. These studies provided the first insight that Polypodiaceae were not monophyletic, because Grammitidaceae appear nested within it. However, in these studies, the family Grammitidaceae was represented by a single species, Micropolypodium okuboi (Yatabe) Hayata, the type of Micropolypodium Hayata.

Using three plastid regions (the two genes rbcL and rps4, and the intergenic spacer rps4-trnS) and expanding the sam-pling of so-called polygrammoid ferns (Polypodiaceae s.str. and grammitids) to 98 selected species, Schneider & al. (2004) obtained similar results, and further recovered Grammitida-ceae as monophyletic but still nested within a larger clade cor-responding to Polypodiaceae s.l. Grammitidaceae, therefore, are now usually referred to informally as the “grammitid ferns”. Based on these findings, Smith & al. (2006, 2008), in their classification for the leptosporangiate ferns, included all genera formerly treated as Grammitidaceae (e.g., Parris, 1990; Smith, 1995) in Polypodiaceae, which, according to this circumscrip-tion, would comprise ca. 1200 species and 56 genera.

In their study of the Hawaiian Adenophorus Gaudich., Ranker & al. (2003) were the first to focus molecular phyloge-

netic analyses (using rbcL, atpB, and trnL-F) on a grammitid fern genus. The first phylogenetic study focusing on the gram-mitid ferns worldwide, however, was conducted by Ranker & al. (2004), who included 73 species of grammitid ferns (rep-resenting 16 genera) and using molecular (plastid sequences atpB and rbcL) as well as morphological characters. Results of this latter study highlighted that many of the traditionally recognized genera were not monophyletic. For example, none of the classically accepted genera, e.g., Ctenopteris Blume, Gram-mitis Sw., Prosaptia C. Presl, and Xiphopteris Kaulf. were supported as monophyletic. Furthermore, some more recently described genera, e.g., Lellingeria A.R. Sm. & R.C. Moran, and Terpsichore A.R. Sm., were also not supported as mono-phyletic.

In the last few years, phylogenetic studies have been per-formed to resolve relationships within predominantly New World grammitids, i.e., Melpomene A.R. Sm. & R.C. Moran (Lehnert & al., 2009), Lellingeria (Labiak & al., 2010a, b), Terpsichore (Sundue & al., 2010), and the “Lellingeria myo-suroides clade” (Ranker & al., 2010).

Ranker & al. (2004) showed that Micropolypodium was not monophyletic, with species of the genus Terpsichore nested within it. In their analyses, out of 24 known Micropolypodium species (Smith, 1992: 22 in the New World and two in the Pa-leotropics; none shared by these two areas), they sampled only three Neotropical species (Micropolypodium taenifolium (Jen-man) A.R. Sm., M. zurquinum (Copel.) A.R. Sm., M. hyalinum

Moranopteris: A new Neotropical genus of grammitid ferns (Polypodiaceae) segregated from Asian Micropolypodium

Regina Y. Hirai,1 Germinal Rouhan,2 Paulo H. Labiak,3 Tom A. Ranker4 & Jefferson Prado1

1 Instituto de Botânica, Caixa Postal 68041, CEP 04045-972, São Paulo-SP, Brazil2 Muséum national d’Histoire naturelle, UMR CNRS 7205 ‘Origine, Structure et Evolution de la Biodiversité’, Botanique,

16 rue Buffon CP 39, 75005 Paris, France3 Universidade Federal do Paraná, Departamento de Botânica, Caixa Postal 19031, CEP 81531-980, Curitiba-PR, Brazil4 Department of Botany, University of Hawaii, 3190 Maile Way, Honolulu, Hawaii 96822, U.S.A.Author for correspondence: Regina Y. Hirai, regina.hirai@gmail.com

Abstract Moranopteris, a new genus of grammitid ferns here described, is segregated from Micropolypodium on the basis of molecular phylogenetic analyses (plastid sequences including atpB, rbcL, and trnL-trnF) and morphological characters. Bayesian and maximum parsimony analyses support Moranopteris as sister clade to a group that includes Micropolypodium s.str. and 11 or 12 other genera, mostly from the Old World. A morphological synapomorphy for the species of Moranopteris is the presence of minute and slightly catenate branched hairs (with or without a setiform branch) and sometimes unbranched hairs occurring together on the stipes, costae, and laminar tissues abaxially. In contrast, Micropolypodium s.str. has only minute, catenate, and unbranched hairs borne on the same parts of the leaves. The new genus Moranopteris comprises 28 spe-cies—including two new species here described, M. rupicola and M. simplex—and one hybrid, occurring in the Neotropics. The 27 necessary new combinations are here proposed. Our results suggest that Micropolypodium is confined to eastern Asia.

Keywords cpDNA; Grammitidaceae; molecular phylogeny; Neotropics; Paleotropics

Supplementary Material The alignment is available in the Supplementary Data section of the online version of this article (http://www.ingentaconntect.com/content/iapt/tax).

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(Maxon) A.R. Sm.), and no Asian taxa. Mickel & Smith (2004) mentioned some preliminary, unpublished molecular results on Neotropical species of Micropolypodium indicating that the Asian species (including the type of the generic name, M. oku-boi) of Micropolypodium s.str. were not most closely related to the Neotropical species.

According to Ranker & al. (2004), the Neotropical spe-cies of Micropolypodium formed a well-supported clade that included two, and possibly three, species of the Terpsichore achilleifolia group as defined by Smith (1993): T. achilleifolia (Kaulf.) A.R. Sm., T. longisetosa (Hook.) A.R. Sm., and, pos-sibly, T. gradata (Baker) A.R. Sm. Although the last taxon was not included in analyses by Ranker & al. (2004), it appeared to be related to the two other species of Terpsichore, and all three species are morphologically similar to Micropolypodium because of their orangish to golden rhizome scales, leaves with determinate growth, conspicuous hydathodes on the adaxial laminar surfaces, and dark red-brown unbranched setae on laminar tissues. Based on these results of Ranker & al. (2004), Labiak & Matos (2007) described and assigned a new hybrid from Rio de Janeiro (Brazil) to Micropolypodium (M. ×bra-dei Labiak & F.B. Matos), and also combined T. achilleifolia and T. gradata in the genus (= M. achilleifolium (Kaulf.) Labiak & F.B. Matos and M. gradatum (Baker) Labiak & F.B. Matos).

Because Ranker & al. (2004) did not include any Paleotrop-ical Micropolypodium in their sampling, and because levels of morphological homoplasy are high among the Neotropical and Paleotropical species of Micropolypodium (and more generally among all grammitid ferns; Ranker & al. 2004), we conducted molecular phylogenetic studies focused on Micropolypodium with a much broader sampling of the species named in the genus. The three major aims of this study were (1) to test the monophyly of Micropolypodium s.l.; (2) to confirm the mono-phyly of the Neotropical species of Micropolypodium, as sug-gested by previous studies (Ranker & al., 2004); and (3) to investigate the evolutionary relationships between Micropoly-podium and other grammitid genera; the here presented results will lay the foundation for a taxonomic revision of the lineage.

MATERIALS AND METHODS

Taxonomic sampling. — Outgroup taxa were selected based on the studies by Schneider & al. (2004), and in-cluded Polypodium vulgare L., Pecluma eurybasis (C. Chr.) M.G. Price, Microgramma bifrons (Hook.) Lellinger, Micro-gramma percussa (Cav.) de la Sota, and Serpocaulon triseriale (Sw.) A.R. Sm.

To test the monophyly of Micropolypodium s.l., and to at-tempt to identify its sister group, representatives of most Neo-tropical and Paleotropical grammitid genera were included in the sampling. They represent all major clades known for grammitids (Ranker & al., 2004).

For Micropolypodium, two out of three known Asian spe-cies and 19 out of 24 known Neotropical species (Smith, 1992; Labiak & Matos, 2007) were sampled, plus Terpsichore longi-setosa. The final matrix included 75 terminals, corresponding

to 62 species representing 22 grammitid genera. Voucher in-formation and GenBank accession numbers are listed in the Appendix.

DNA extraction. — Genomic DNA was isolated from ei-ther field-collected, silica gel–dried leaves or from herbarium specimens when fresh material was unavailable. Total genomic DNA was extracted using the Qiagen DNeasy Plant Mini Kit (Valencia, California, U.S.A.) following the manufacturer’s protocol, but with an additional proteinase K digestion during the lysis step for the herbarium material: 30 μL of proteinase K (20 mg/mL), and 30 μL of β-mercaptoethanol (98%) were added per tube, and the tubes were incubated on a tipping plate at 42°C for 12 hours.

Amplifications and sequencing. — Three plastid DNA sequences were amplified by PCR, the two coding regions atpB and rbcL, and the non-coding spacer trnL-trnF. For atpB, we used the primers ESATPB172F and ESATPE45R (Schuettpelz & Pryer, 2007), for rbcL the primers ESRBCL1F and ESRB-CL1361R (Schuettpelz & Pryer, 2007), and for trnL-trnF the primers “e” and “f  ” (Taberlet & al., 1991). The two genes were typically amplified using a program beginning with one initial denaturation step of 5 min at 94°C, followed by 35 cycles of 1 min at 94°C, 1 min at 50°C, 2.5 min at 72°C, and a final extension period of 10 min at 72°C. For trnL-F, we used the following protocol: 5 min at 94°C/35× (1 min, 94°C; 30 s, 50°C; 1 min, 72°C)/7 min at 72°C. The PCR products were sequenced by the High-Throughput Genomics Unit at the University of Washington, using the amplification primers plus the internal primers 493F and 910R for atpB (Schuettpelz & Pryer, 2007), and ESRBCL628F and ESRBCL654R for rbcL (Schuettpelz & Pryer, 2007).

Alignment and phylogenetic analyses. — Forward and reverse sequences obtained for all species were edited and as-sembled using Sequencher v.4 (Gene Codes Corporation, Ann Arbor, Michigan, U.S.A.). Consensus sequences were then au-tomatically aligned using Muscle v.3.6 using the default param-eters (Edgar, 2004), and the resulting alignments were manu-ally checked using the MUST package (Philippe, 1993) and revised to maximize the similarity between sequences. Data matrices were constructed using TaxonDNA v.1.6.2. (Meier & al., 2006) and analyzed using equally weighted maximum parsimony and Bayesian inference, with indels treated as miss-ing data.

Maximum parsimony analyses (MP) were performed us-ing the program PAUP* v.4.0b10 (Swofford, 2002). For all MP analyses, heuristic searches were performed with 10,000 ran-dom-sequence addition replicates, tree bisection-reconnection (TBR), branch swapping, and the Multrees options were on. The robustness of each node was assessed by a non-parametric bootstrap analysis (BS; Felsenstein, 1985), with 1000 replicates of similar heuristic searches (each replicate with 3 random-se-quence addition replicates). Because all characters were equally sampled, parsimony-uninformative characters could have had a significant effect on robustness and were removed before the bootstrap procedure (Desalle & al., 2002). For comparison, another bootstrap procedure was conducted leaving all char-acters including parsimony-uninformative ones; the resulting

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bootstrap values were unchanged or very similar with respect to each node (values differed by at most 4% from a procedure to the other one).

Bayesian analyses (BI) were performed using the program MrBayes v.3.1.2 (Huelsenbeck & Ronquist, 2001; Ronquist & Huelsenbeck, 2003). The GTR + Γ + I model of nucleotide substitution was identified as the best fit to the data for each of the three regions under the Akaike information criterion (AIC; Akaike, 1973) as implemented in the program jModelTest v.0.1.1 (Posada, 2008). Two independent but parallel analy-ses were conducted using flat priors, starting from random trees and consisting of four chains each. The analyses were run for 5 million generations, sampling every 100 generations. Following completion of the analyses, the output parameter estimates through time were explored in order to recognize the point of convergence to the stationary distribution, using Tracer v.1.5 (Rambaut & Drummond, 2004). This was reached after approximately 150,000 generations, and the first 500,000 generations (5000 trees) of one of the runs were conservatively excluded as the “burn-in”. We pooled the post–burn-in trees (45,000 total trees) and computed a 50% majority-rule consen-sus with average branch lengths and posterior probability (PP) estimates for all nodes.

Taxonomic treatment. — For the Neotropical Micropoly-podium clade, which is here described as a new genus, all the necessary new combinations are proposed, as well as a com-plete list of homotypic synonyms for each species. Based on morphology, two new species are described. In particular, spores and laminar tissues harbor critical characters (the micro-morphology of the perispores and bases of setae, respectively). Taken from herbarium specimens of Micropolypodium at NY and UC, spores and laminar tissues were imaged using a scan-ning electron microscope (SEM). Spores and laminar tissues were transferred with dissecting needles from the specimens to SEM stubs coated with an adhesive tape. The stubs were then coated with gold-palladium in a sputter-coater for 30 to 40 seconds and imaged digitally using a JEOL JSM-5410LV SEM equipped with a JEOL Orion 5410 software interface. The accelerating voltage was 15 kV.

Leaf tissue clearings were made for almost all species, using the Strittmatter method (1973; apud Kraus & Arduin,

1997). These leaves were drawn to show the venation patterns and hairs in the genus. Hairs were measured on Herbarium specimens.

RESULTS

Results for each analysis and for each DNA region, includ-ing the number of characters and taxa included, percentages of variable and informative sites, number of MP trees, tree lengths, consistency and retention indices (CI; RI), and substi-tution models used in BI are summarized in Table 1.

The analyses performed for the combined dataset produced trees with improved resolution and higher support node values compared to those obtained from separate datasets. Thus, only the results from the combined dataset using MP and BI analy-ses are described and discussed. Tree topologies were similar regardless of the analytical method employed, i.e., the strict consensus obtained in MP was nearly identical to the majority consensus tree resulting from BI analyses, except for some unresolved relationships and the placement of three terminals, Chrysogrammitis Parris, Ctenopterella Parris, and Grammitis deplanchei (Baker) Copel. (Fig. 1).

The results of these analyses show Micropolypodium s.l. as not monophyletic. All Neotropical species of Micropoly-podium s.l. form a clade (M. truncicola (Klotzsch) A.R. Sm. through M. grisebachii (Underw. ex C. Chr.) A.R. Sm.) that is strongly supported (Fig. 1; BS = 94%; PP = 1.0). The two Asian species of Micropolypodium are grouped together (M. okuboi and M. sikkimense (Hieron.) X.C. Zhang; BS = 78%; PP = 1.0) and rooted in a broader clade comprising 11 other Paleotropical grammitid genera in BI (Calymmodon C. Presl, Scleroglossum Alderw., Tomophyllum (E. Fourn.) Parris, Dasygrammitis Par-ris, Ctenopterella, Grammitis, Oreogrammitis Copel., Radio-grammitis Parris, Themelium (T. Moore) Parris, Prosaptia, Ctenopteris) or 12 in MP analyses (adding Chrysogrammitis).

Within the Neotropical clade, here treated as the new ge-nus Moranopteris, the basal polytomy resulting from the MP analyses precludes the assessment of the relationships among the species. Although this polytomy is resolved in BI, the re-lationships are poorly supported (PP < 0.9). Two major sister

Table 1. Number of taxa and characters, evolutionary models used in BI, and tree statistics for the MP analyses.atpB rbcL trnL-trnF Combined

Total number of taxa/ingroup only 69/31 71/33 57/32 75/36Included characters 1357 1311 404 3072Variable characters (%) 359 (26) 346 (26) 259 (64) 964 (31)Parsimony-informative characters (%) 254 (19) 236 (18) 192 (47) 682 (22)Consistency index 0.45 0.43 0.50 0.44Retention index 0.65 0.65 0.70 0.64Number of MP trees 156 56,243 458 12MP tree length 979 987 845 2938BI substitution model GTR + Г + I GTR + Г + I GTR + Г + I GTR + Г + I

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Fig. 1. Concatenated phylogram obtained with the Bayesian inference and strict consensus of the 12 most parsimonious trees resulting from the MP analysis, based on the combined dataset atpB + rbcL + trnL-F. Irresolution and topology differing in the strict consensus MP tree are indicated by dashed lines. Statistical support values PP > 0.50 and BS > 50 are indicated on the branches as PP/BS, with PP = 1 and BS = 100% appearing as “+”. Geographic distributions are indicated in the grey frames for the two distinct clades discussed in the text, the new genus Moranopteris and Micropolypodium s.str.

+/72

.92/-

+/+

+/99

+/96

+/75

+/94

.95/46

+/75

+/75

+/91

+/95 Moranopteris truncicolaMoranopteris truncicola

+/+ Moranopteris taenifoliaMoranopteris taenifolia

+/79+/96

+/+ Moranopteris caucanaMoranopteris caucana

Moranopteris cookiiMoranopteris williamsii

+/91+/+ Moranopteris microlepis

Moranopteris microlepisMoranopteris plicata

+/98

+/+.99/- Moranopteris serricula

Moranopteris serriculaMoranopteris zurquina

+/+ Moranopteris trichomanoidesMoranopteris trichomanoides

+/84+/+

.99/76 Moranopteris blepharideaMoranopteris hyalinaMoranopteris hyalinaMoranopteris hyalinaMoranopteris perpusillaMoranopteris nanaMoranopteris setosa

Moranopteris achilleifoliaMoranopteris gradata

+/64

+/+.98/79 Moranopteris aphelolepis

Moranopteris aphelolepisMoranopteris basiattenuata

+/92 Moranopteris longisetosaMoranopteris longisetosa

Moranopteris grisebachii

.96/57

+/-

+/96

+/70+/75 Oreogrammitis hookeri

Radiogrammitis parvaThemelium conjunctisorum

+/51 Prosaptia contiguaProsaptia nutans

Ctenopterella denticulata

.92 /-

+/97+/84 Grammitis poeppigiana

Grammitis billardiereiCtenopteris heterophyllaGrammitis deplanchei

Ctenopteris lasiostipes

+/-

+ /56

+/78+/75

+/93 Micropolypodium okuboiMicropolypodium okuboi

Micropolypodium okuboi+/+ Micropolypodium sikkimense

Micropolypodium sikkimenseCalymmodon gracilis

+/69 Scleroglossum sulcatumTomophyllum repandulum

Dasygrammitis crassifrons+/+ Chrysogrammitis glandulosa

Chrysogrammitis musgraviana

.99

+/99Lellingeria apiculata

Lellingeria limula+/+ Melpomene moniliformis

Melpomene flabelliformisAscogrammitis pichinchae

+/96 Enterosora percrassaCeradenia spixiana

11 Terpsichore lanigera

Terpsichore cultrataLeucotrichum schenckii

+/+Cochlidium punctatum

Cochlidium serrulatumGrammitis bryophila

Adenophorus tripinnatifidusGrammitis tenellaTerpsichore lehmannianaTerpsichore asplenifolia

Serpocaulon triseriale+/65 Microgramma percussa

Microgramma bifronsPolypodium vulgare

Pecluma eurybasis0.1 substitution/site

-/63

.61/67

-/73

+/+

+/+

+/+

-/66

Moranopteris(formerly: Micropolypodium s.l.)

Micropolypodium s.str.

Paleotropics

Neotrop

ics

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clades were well supported, the first comprising M. blephari-dea and M. hyalina (4 terminals BS = 84%; PP = 1.0), and the second consisting of 10 species from M. truncicola to M. tri-chomanoides (16 terminals, BS = 75%; PP = 1.0).

The MP analysis shows a large, poorly supported clade (BS = 57%) formed of primarily Paleotropical species of grammitid genera (including Micropolypodium s.str. and Chrysogrammi-tis) as the sister group to the Neotropical Moranopteris clade. In the BI analysis, Chrysogrammitis from SE Asia through Male-sia to Melanesia (Parris, 1998) is recovered as sister group to the large clade (PP = 1.0) comprising the Neotropical Moranopteris clade plus the clade with Micropolypodium s.str. and other Paleotropical genera.

The relationships among Paleotropical genera remain un-clear with both analyses. Calymmodon is weakly supported (BS < 70%; PP = 0.61) as sister to Micropolypodium s.str. Addition-ally, Scleroglossum and Tomophyllum form a clade that is closely related to Micropolypodium s.str. plus Calymmodon (BS = 56%; PP = 1.0).

DISCUSSION

Our results support two distinct clades of Micropolypo-dium. One includes all of the sampled Neotropical species (Moranopteris) and the other all of the sampled Paleotropical species, as previously suggested by Ranker & al. (2004) and Mickel & Smith (2004).

The Neotropical species of Micropolypodium differ from the Paleotropical species of Micropolypodium most obviously in the morphology of the hairs on fronds. They are slightly cat-enate, unbranched and branched on stipes, rachises, or costae and laminar tissues, mainly abaxially, whereas hairs are only catenate and unbranched on the Paleotropical taxa. There may be other characters that are exclusive to the Neotropical clade of Micropolypodium s.l., but additional studies are necessary to address this issue (e.g., the ontogeny of setae and leaf vena-tion patterns).

Our results corroborate the placement of the three species considered in the Terpsichore achilleifolia group (group 5 of Smith, 1993) as being members of the clade with the Neotropi-cal species of Micropolypodium, as first suggested by Ranker & al. (2004) and supported by Sundue & al. (2010). In addition to the molecular phylogenetic support, taxa of the T. achillei-folia group are morphologically more similar to Neotropical Micropolypodium than to the remaining groups of Terpsichore s.l., from which they differ by displaying orangish to golden rhizome scales and red-brown unbranched setae (Smith, 1993).

Parris (2009) recognized Micropolypodium s.str. in Male-sia as having radial rhizomes with fronds in whorls, and as having pale to medium brown, non-clathrate and glabrous rhi-zome scales, laminae deeply pinnately divided to pinnate, one sorus per lobe of pinna, medium to dark brown simple hairs (i.e., no branched hairs), hydathodes, and glabrous sporangia. The Neotropical species of Micropolypodium (= Moranopteris) share most of those features with the Paleotropical taxa except that they have branched hairs.

Calymmodon is putatively sister to Micropolypodium s.str. Furthermore, Scleroglossum plus Tomophyllum form a mono-phyletic group, sister to Micropolypodium s.str. plus Calym-modon; although not strongly supported, these relationships were found in previous studies (Ranker & al., 2004; clade III). The placement of Chrysogrammitis is still uncertain, as found by Ranker & al. (2004). It is strongly supported only with BI, recovered as the sister clade to Paleotropical grammitid genera plus the Neotropical clade of Micropolypodium s.l. (includ-ing the Terpsichore achilleifolia group). By contrast in MP analyses, Chrysogrammitis appeared as a weakly supported sister group to Ctenopterella and Grammitis, both included in the Paleotropical clade. Schuettpelz & Pryer (2007), in their phylogenetic study based on three plastid genes (rbcL, atpB, atpA), retrieved the Paleotropical Chrysogrammitis musgra-viana (Baker) Parris as sister group to Micropolypodium s.l. (M. hyalinum and M. taenifolium), although it was not strongly supported either. Sundue & al. (2010) obtained a similar result, i.e., the Neotropical species of Micropolypodium (including Terpsichore longisetosa) appeared in a strongly supported clade (BS = 100%), and this clade is sister to Chrysogram-mitis musgraviana.

Based on phylogenetic evidence, as well as morphological characters, the Neotropical species of Micropolypodium s.l. are here segregated from Micropolypodium s.str., a name now restricted to a group from eastern Asia. We propose a new generic name Moranopteris for the Neotropical clade, with the following description:

Moranopteris R.Y. Hirai & J. Prado, gen. nov. – Type: Mo-ranopteris basiattenuata (Jenman) R.Y. Hirai & J. Prado (see below).Diagnosis. – Primo aspectu ad Micropolypodio valde ac-

cedit, sed frondibus cum pilis leviter catenulatis ramosis et non-ramosis, pilis ramosis cum ramis setaceis et non-setaceis (vs. frondibus solum cum pilis non-ramosis), distributio neo-tropica (vs. veteri-mundi) distincta.

Description. – Plants epiphytic, rarely epipetric or ter-restrial. Rhizomes short-creeping, decumbent to ascending, slightly dorsiventral to radially symmetric, scaly, the scales golden to castaneous, often lustrous, nonclathrate, the cells of the scales body turgid and shining, margins entire or bear-ing small, hyaline, lateral projections, or with glanduliform or setiform projections, these setiform projections mostly red-brown, rarely hyaline, apical or subapical gland-like cells pres-ent or not. Leaves monomorphic, caespitose, erect to pendent, determinate, or rarely ± indeterminate (M. caucana, M. kil-lipii), short-stipitate, sometimes the stipes almost absent, not articulate to the rhizomes; laminae mostly linear, sometimes lanceolate, pinnatifid or pinnatisect, rarely bipinnatisect or pinnate-pinnatisect, sometimes pinnate only at base, gradually tapering proximally to a lateral narrow wing or not, mostly chartaceous, sometimes membranaceous; costae obscured by greenish laminar tissue or overlain by dark sclerenchyma ab-axially and adaxially; segments ascending to patent, alternate to opposite, with an acroscopic hump (segments gibbous) or not, especially in fertile segments, entire, oblong or rounded,

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rarely linear and pinnatisect or pinnatifid (M. achilleifolia), slightly decurrent at base or not, margins plane or slightly revo-lute, sometimes hyaline marginally, basiscopic side not folded over sorus; veins simple or furcate, rarely pinnate, inconspicu-ous, obscured by the laminar tissue or by dark sclerenchyma at the bases of segments, hydathodes present, mostly 1 or 2 (rarely > 2) per segment or pinna, visible adaxially, elliptic, round, or oblong; setae dark red-brown, present on both sur-faces of the stipes, rachises, and/or costae, also sometimes on laminar tissue, rarely absent, ring of cells at base of setae conspicuous to inconspicuous; hairs present on the stipes, ra-chises, and/or costae, also on laminar tissue (mainly near costae abaxially), hyaline, slightly red-brown, mostly branched, or unbranched and branched. Sori superficial, usually at base of the costae/costules or on the acroscopic veinlet, mostly 1 (rarely > 2) per segment or pinna, rounded or elliptic, exindusiate; soral paraphyses absent; sporangia glabrous; spores tetrahedral-globose, papillate to tuberculate with small globules; x = 37 (Smith, 1992).

Distribution. – Moranopteris occurs in the Neotropics, with 28 known species and one hybrid. It occurs in Mexico, Central America, Antilles, Andes, southern Venezuela, Guyanas, and Brazil. Andes and Central America are centers of diversity of the genus, with 14 and 12 species, respectively. In these centers, five species are endemic to the Andes (Moranopteris aphelol-epis, M. blepharidea, M. killipii, M. simplex, M. williamsii), and one species to Central America (M. zurquina). Three species and one hybrid are endemic to Brazil (M. gradata, M. per-pusilla, M. setosa, M. ×bradei) and two species are endemic to the Caribbean region (M. knowltoniorum, M. sherringii). Moranopteris nimbata, primarily a Caribbean species, was also reported by Farrar (1967) for North Carolina. It is the unique record for Moranopteris outside of the Neotropics.

Distinctive morphological characters. – Moranopteris dif-fers from Micropolypodium s.str. in having slightly catenate unbranched and branched hairs intermixed together on the same fronds (Fig. 2L–P) vs. only catenate unbranched hairs (similar to Fig. 2L).

The rhizome scales are nonclathrate and the cells of the scale body are often turgid and shining. The margins of rhi-zome scales are mostly entire, or with small lateral projections, or with setiform or glanduliform projections (Fig. 2G–K).

Blade segments sometimes are gibbous, mainly on fertile segments (Fig. 2F). The number of hydathodes and veins is 1 or 2 (Fig. 2E, F), or sometimes more than two per segment (Moranopteris achilleifolia; M. gradata, Fig. 2C; M. longise-tosa, Fig. 2D).

Setae are commonly present on the stipes, rachises, cos-tae, and laminar tissue, but are absent in Moranopteris grise-bachii and M. perpusilla. As noted by Smith (1992), setae in the Neotropical species of Micropolypodium look different from those of many other grammitid ferns. He described the setae as bearing a ring of cells at their bases. These cells are different from the adjacent surrounding epidermal cells, as in Moranopteris basiattenuata (Fig. 3A) and in several other spe-cies examined. Moreover, Smith (pers. comm.) observed that Micropoly podium s.str. species do not have this ring of cells,

such as in Micropolypodium okuboi (Fig. 3B). We observed this ring of cells at the bases of setae in many species of Moranop-teris, but it is inconspicuous in some plants, and it is a difficult character to use to quickly identify members of the genus.

Hairs are often 1–4-branched, and sometimes there are un-branched hairs and branched hairs occurring on the same plant (Fig. 2L–P). The branched hairs can have 1 (Fig. 2P) or some-times 2 setiform branches. Some species (e.g., Moranopteris blepharidea and M. killipii) can be recognized by 1-branched hairs that typically have a longer branch, at least twice as long as the main branch (Fig. 2M).

Spores of Moranopteris appear to be only slightly differ-ent from those of Micropolypodium. Tryon & Lugardon (1990) described the spores of Grammitis knowltoniorum (= Moranop-teris knowltoniorum) as being tuberculate. Perispores of Mo-ranopteris basiattenuata also display tuberculate surfaces with spherical globules (Fig. 3C, D), whereas they are papillate with dense globules in Micropolypodium okuboi (Fig. 3E, F), and the surface is depressed with only low papillae in spores of Micropolypodium sikkimense (Fig. 3G, H).

Etymology. – The genus is named in honor of Dr. Robbin C. Moran, fern specialist and Curator of ferns and ly-cophytes at The New York Botanical Garden (NY).

New combinations

1. Moranopteris achilleifolia (Kaulf.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium achilleifolium Kaulf., Enum. Filic.: 116. 1824 ≡ Ctenopteris achilleifolia (Kaulf.) J. Sm., Hist. Fil.: 185. 1875 ≡ Grammitis achilleifolia (Kaulf.) R.M. Tryon & A.F. Tryon in Rhodora 84: 128. 1982 ≡ Terp-sichore achilleifolia (Kaulf.) A.R. Sm. in Novon 3: 486. 1993 ≡ Micropolypodium achilleifolium (Kaulf.) Labiak & F.B. Matos in Brittonia 59: 184. 2007. – Distribution: Coastal Brazil and probably also in Argentina (Missiones).

2. Moranopteris aphelolepis (C.V. Morton) R.Y. Hirai & J. Prado, comb. nov. ≡ Grammitis aphelolepis C.V. Mor-ton in Contr. U.S. Natl. Herb. 38: 97. 1967 ≡ Xiphopteris aphelolepis (C.V. Morton) Pic.Serm. in Webbia 28: 472. 1973 ≡ Micropolypodium aphelolepis (C.V. Morton) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Colombia, Ecuador, and Bolivia.

3. Moranopteris basiattenuata (Jenman) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium basiattenuatum Jenman in Bull. Bot. Dept. Jamaica, n.s., 4: 114. 1897 ≡ Xiphopteris basiat-tenuata (Jenman) Copel. in Amer. Fern J. 42: 104. 1952 ≡ Grammitis basiattenuata (Jenman) Proctor in Bull. Inst. Ja-maica, Sci. Ser. 5: 32. 1953 ≡ Micropolypodium basiattenu-atum (Jenman) A.R. Sm. in Novon 2: 422. 1992 – Lecto-type first step (Proctor, 1985: 575: “Jenman s.n. (NY)”) and second step (here designated): Jamaica, without exact locality, s.d., G.S. Jenman s.n. (NY-127117!); isolectotype: NY s.n.! – Distribution: Mexico, Guatemala, Honduras, El Salvador, Costa Rica, Jamaica, Dominican Republic, Guyana, Venezuela, Colombia, and Ecuador.

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Fig. 2. Morphology of some species of Moranopteris. A–D, Venation patterns (cleared leaf): A, simple veins, M. cookii (Sundue & al. 1764, NY); B, furcate veins, M. aphelolepis (Boeke & Jaramillo 2701, NY); C, pinnate veins, M. gradata (Hirai & al. 564, SP); D, pinnate veins, M. longiseto-sa (Rojas 3209, NY); E–F, segments: E, not gibbous segments (arrow indicates adaxial hydathode), M. cookii (Sundue & al. 1764, NY); F, gibbous segments at circle (arrows indicate adaxial hydathodes), M. liesneri (Cowan & Wurdack 31415, NY); G–K, rhizome scale types: G, entire margins, M. cookii (Sundue & al. 1764, NY); H, margins with small lateral projections, M. basiattenuata (Breedlove & Thorne 30160, NY); I, margins with glanduliform projections, M. longisetosa (Rojas 3209, NY); J, margins with short setiform projections, M. killipii (Lehmann 7655, K); K, margins with setiform projections, M. microlepis (Rojas 3603, NY); L–P, some hair types (re-hydrated hairs): L, unbranched hairs, M. liesneri (Cowan & Wurdack 31415, NY); M, 1-branched, M. blepharidea (Kessler & al. 11973, UPCB); N, 2-branched, M. longisetosa (Rojas 3209, NY); O, 4-branched, M. setosa (Hirai & al. 598, SP); P, 1-branched with one setiform branch, M. taenifolia (Granville 3557, CAY).

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Fig. 3. A, Base of a seta showing a ring of cells in Moranopteris basiattenuata (Maxon 9858, NY); B, base of a seta without a ring of cells in Micropolypodium okuboi (Togashi s.n., UC); C–D, spores of M. basiattenuata (Breedlove & Thorne 30160, NY): C, proximal face with spherical globules; D, detail of tuberculate surface; E–F, spores of M. okuboi (Togashi s.n., UC): E, proximal face depressed and densely globulose; F, detail of globules on a irregular surface; G–H, spores of Micropolypodium sikkimense (Miehe 93-32, UC): G, proximal face; H, detail of low pappilae on the surface. A–C, E, G: scale bars = 10 μm; D, F, H: scale bars = 2 μm.

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4. Moranopteris blepharidea (Copel.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium blepharideum Copel. in Univ. Calif. Publ. Bot. 19: 304. pl. 64. 1941 ≡ Xiphopteris blepha-ridea (Copel.) Copel. in Amer. Fern J. 42: 99. 1952 ≡ Micro-polypodium blepharideum (Copel.) A.R. Sm. in Novon 2: 422. 1992 ≡ Grammitis blepharidea (Copel.) Stolze in Fieldiana, Bot., n.s., 32: 112. 1993. – Distribution: Colom-bia, Peru, and Bolivia.

5. Moranopteris caucana (Hieron.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium caucanum Hieron. in Bot. Jahrb. Syst. 34: 503. 1904 ≡ Xiphopteris caucana (Hieron.) Copel. in Amer. Fern J. 42: 98. 1952 ≡ Grammitis caucana (Hieron.) C.V. Morton in Contr. U.S. Natl. Herb. 38: 96. 1967 ≡ Micropolypodium caucanum (Hieron.) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Nicaragua, Costa Rica, Panama, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia, and Brazil.

6. Moranopteris cookii (Underw. & Maxon) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium cookii Underw. & Maxon in Contr. U.S. Natl. Herb. 17: 408. 1914 ≡ Xiphop-teris cookii (Underw. & Maxon) Copel. in Amer. Fern J. 42: 98. 1952 ≡ Grammitis cookii (Underw. & Maxon) F. Seym. in Phytologia 31: 173. 1975 ≡ Micropolypodium cookii (Underw. & Maxon) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Guatemala, Costa Rica, Panama, and Ec-uador.

7. Moranopteris gradata (Baker) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium gradatum Baker in Martius, Fl. Bras. 1(2): 513. 1870 ≡ Polypodium hirsutulum Fée, Crypt. Vasc. Brésil 1: 87. 1869, nom. illeg. (ICBN Art. 53.1), non G. Forst. (1786) ≡ Ctenopteris gradata (Baker) Copel. in Philipp. J. Sci. 84: 437. 1955 ≡ Grammitis gradata (Baker) R.M. Tryon & A.F. Tryon in Rhodora 84: 128. 1982 ≡ Terpsichore gradata (Baker) A.R. Sm. in Novon 3: 486. 1993 ≡ Micropolypodium gradatum (Baker) Labiak & F.B. Matos in Brittonia 59: 184. 2007. – Distribution: Endemic to Coastal Brazil.

8. Moranopteris grisebachii (Underw. ex C. Chr.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium grisebachii Underw. ex C. Chr., Index Filic.: 531. 1906, based on P. exiguum Griseb., non Heward (1838) ≡ Polypodium exiguum Griseb., Fl. Br. W. Ind.: 701. 1864, nom. illeg. (ICBN Art. 53.1), non Heward (1838) ≡ Xiphopteris grisebachii (Un-derw. ex C. Chr.) Copel. in Amer. Fern J. 42: 95. 1952 ≡ Grammitis grisebachii (Underw. ex C. Chr.) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 33, t. 2(7–8). 1953 ≡ Mi-cropolypodium grisebachii (Underw. ex C. Chr.) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Endemic to the Greater Antilles and the Lesser Antilles.

9. Moranopteris hyalina (Maxon) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium hyalinum Maxon in Contr. U.S. Natl. Herb. 17: 406. 1914 ≡ Xiphopteris hyalina

(Maxon) Copel. in Amer. Fern J. 42: 107. 1952 ≡ Gram-mitis hyalina (Maxon) F. Seym. in Phytologia 31: 174. 1975 ≡ Micropolypodium hyalinum (Maxon) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Costa Rica, Colombia, Ecuador, and Peru.

10. Moranopteris killipii (Copel.) R.Y. Hirai & J. Prado, comb. nov. ≡ Xiphopteris killipii Copel. in Amer. Fern J. 42: 105, t. 10. 1952 ≡ Grammitis killipii (Copel.) Lellinger in Amer. Fern J. 74: 58. 1984. – Distribution: Endemic to Andes from Colombia and Ecuador.

11. Moranopteris knowltoniorum (Hodge) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium knowltoniorum Hodge in Amer. Fern J. 31: 105, p. 1, f. 4–6. 1941 ≡ Xiphopteris knowlto-niorum (Hodge) Copel. in Amer. Fern J. 42: 108. 1952 ≡ Grammitis knowltoniorum (Hodge) Proctor in Rhodora 63: 35. 1961 ≡ Micropolypodium knowltoniorum (Hodge) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Endemic to the Lesser Antilles.

12. Moranopteris liesneri (A.R. Sm.) R.Y. Hirai & J. Prado, comb. nov. ≡ Grammitis liesneri A.R. Sm. in Ann. Mis-souri Bot. Gard. 77: 257. 1990 ≡ Micropolypodium liesneri (A.R. Sm.) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Endemic to Venezuela.

13. Moranopteris longisetosa (Hook.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium longisetosum Hook., Sp. Fil. 4: 225, t. 278 A. 1864 ≡ Terpsichore longisetosa (Hook.) A.R. Sm. in Novon 3: 487. 1993. – Distribution: Costa Rica, Venezuela, Colombia, Ecuador, Peru, and Bolivia.

14. Moranopteris microlepis (Rosenst.) R.Y. Hirai & J. Prado, comb. et stat. nov. ≡ Polypodium blepharodes Maxon var. microlepis Rosenst. in Repert. Spec. Nov. Regni Veg. 22: 14. 1925. – Distribution: Costa Rica, Panama, Jamaica, and Guadeloupe.

15. Moranopteris nana (Fée) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium nanum Fée, Mém Foug. 5, Gen. Filic.: 238. 1852 ≡ Xiphopteris nana (Fée) Copel. in Amer. Fern J. 42: 107. 1952 ≡ Micropolypodium nanum (Fée) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Costa Rica, Panama, Trinidad and Tobago (restricted to the island of Trinidad), French Guiana, Suriname, Guyana, Venezuela, Colombia, Peru, and Brazil.

16. Moranopteris nimbata (Jenman) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium nimbatum Jenman in J. Bot. 24: 271. 1886 ≡ Xiphopteris nimbata (Jenman) Copel. in Amer. Fern J. 42: 108. 1952 ≡ Grammitis nimbata (Jenman) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 34. 1953 ≡ Micro-polypodium nimbatum (Jenman) A.R. Sm. in Novon 2: 422. 1992. – Distribution: U.S.A. (North Carolina; Farrar, 1967) and the Greater Antilles.

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17. Moranopteris perpusilla (Maxon) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium perpusillum Maxon in Contr. U.S. Natl. Herb. 17: 409, pl. 13-A. 1913 ≡ Xiphopteris per-pusilla (Maxon) Copel. in Amer. Fern J. 42: 95. 1952 ≡ Mi-cropolypodium perpusillum (Maxon) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Endemic to Southeast Brazil.

18. Moranopteris plicata (A.R. Sm.) R.Y. Hirai & J. Prado, comb. nov. ≡ Grammitis plicata A.R. Sm. in Ann. Mis-souri Bot. Gard. 77: 258. 1990 ≡ Micropolypodium plica-tum (A.R. Sm.) A.R. Sm. in Novon 2: 422. 1992. – Distri-bution: Costa Rica, Venezuela, Colombia, Ecuador, Peru, and Brazil.

19. Moranopteris serricula (Fée) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium serricula Fée, Mém. Foug. 5, Gen. Filic.: 238. 1852 ≡ Xiphopteris serricula (Fée) Copel. in Amer. Fern J. 42: 101. 1952 ≡ Grammitis serricula (Fée) Proctor in Rhodora 63: 35. 1961 ≡ Micropolypodium ser-ricula (Fée) A.R. Sm. in Novon 2: 422. 1992. – Distribu-tion: Dominica, Guadeloupe, Martinique, Trinidad and To-bago (restricted to the island of Trinidad), and Venezuela.

20. Moranopteris setosa (Kaulf.) R.Y. Hirai & J. Prado, comb. nov. ≡ Xiphopteris setosa Kaulf., Enum. Filic.: 275. 1824 ≡ Grammitis setosa (Kaulf.) C. Presl, Tent. Pterid.: 208. 1836, nom. illeg. (ICBN Art. 53.1), non Blume (1828) ≡ Polypodium micropteris C. Chr., Index Filic.: 545. 1906, nom. nov. for X. setosa Kaulf. ≡ Grammitis micropteris (C. Chr.) Brade in Sellowia 18: 81. 1966 ≡ Micropolypo-dium setosum (Kaulf.) A.R. Sm. in Novon 2: 422. 1992. – Distribution: Endemic to Coastal Brazil.

21. Moranopteris sherringii (Baker) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium sherringii Baker in J. Bot. 20: 26. 1882 ≡ Xiphopteris sherringii (Baker) Copel. in Amer. Fern J. 42: 104. 1952 ≡ Grammitis sherringii (Baker) Proc-tor in Bull. Inst. Jamaica, Sci. Ser. 5: 35. 1953 ≡ Micropoly-podium sherringii (Baker) A.R. Sm. in Novon 2: 423. 1992. – Distribution: Endemic to the Greater Antilles.

22. Moranopteris taenifolia (Jenman) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium taenifolium Jenman in Bull. Bot. Dept. Jamaica, n.s., 4: 114. 1897 ≡ Xiphopteris taenifolia (Jenman) Copel. in Amer. Fern J. 42: 109. 1952 ≡ Grammi-tis taenifolia (Jenman) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 35. 1953 ≡ Micropolypodium taenifolium (Jenman) A.R. Sm. in Novon 2: 423. 1992. – Distribution: Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Pa-nama, Jamaica, Haiti, Dominican Republic, Puerto Rico, Saint Kitts and Nevis, Guadeloupe, Martinique, Trinidad and Tobago (restricted to the island of Trinidad), French Guiana, Suriname, Guyana, Venezuela, Colombia, Ecua-dor, Peru, Bolivia, and Brazil.

23. Moranopteris trichomanoides (Sw.) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium trichomanoides Sw., Prodr.: 131.

1788 ≡ Grammitis trichomanoides (Sw.) Ching in Bull. Fan Mem. Inst. Biol. 10: 16. 1940 ≡ Xiphopteris trichomanoi-des (Sw.) Copel., Gen. Fil.: 215. 1947 ≡ Micropolypodium trichomanoides (Sw.) A.R. Sm. in Novon 2: 423. 1992. – Distribution: Mexico, Guatemala, Honduras, Cuba, Ja-maica, Haiti, and Dominican Republic.

24. Moranopteris truncicola (Klotzsch) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium truncicola Klotzsch var. major Klotzsch in Linnaea 20: 374. 1847 ≡ Xiphopteris trunci-cola (Klotzsch) Copel. in Amer. Fern J. 42: 101. 1952 ≡ Grammitis truncicola (Klotzsch) C.V. Morton in Contr. U.S. Natl. Herb. 38: 98. 1967 ≡ Micropolypodium truncic-ola (Klotzsch) A.R. Sm. in Novon 2: 423. 1992. – Distribu-tion: Costa Rica, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia, and Brazil.

25. Moranopteris williamsii (Maxon) R.Y. Hirai & J. Prado, comb. nov. ≡ Polypodium williamsii Maxon in Contr. U.S. Natl. Herb. 17: 547, t. 34. 1916 ≡ Xiphopteris williamsii (Maxon) Copel. in Amer. Fern J. 42: 101. 1952 ≡ Grammitis williamsii (Maxon) Lellinger in Amer. Fern J. 74: 59. 1984 ≡ Micropolypodium williamsii (Maxon) A.R. Sm. in Novon 2: 423. 1992. – Distribution: Endemic to Bolivia.

26. Moranopteris zurquina (Copel.) R.Y. Hirai & J. Prado, comb. nov. ≡ Xiphopteris zurquina Copel. in Amer. Fern J. 42: 99. 1952 ≡ Grammitis zurquina (Copel.) F. Seym. in Phytologia 31: 175. 1975 ≡ Micropolypodium zurquinum (Copel.) A.R. Sm. in Novon 2: 423. 1992. – Distribution: Endemic to Costa Rica.

27. Moranopteris ×bradei (Labiak & F.B. Matos) R.Y. Hirai & J. Prado, comb. nov. ≡ Micropolypodium ×bradei Labiak & F.B. Matos in Brittonia 59: 182. 2007. – Distribution: Known only from Rio de Janeiro State, southeastern Brazil.

New species

1. Moranopteris rupicola R.Y. Hirai & J. Prado, sp. nov. – Type: GUYANA. Mazaruni-Potaro, Roraima, summit, La Proa Camp, east of the border, near Lake Gladys, 2800 m, 05°15′36″ N, 60°13′ W, 14 Apr 1988, R.L. Liesner 23289 (holotype: MO; isotypes: UC, VEN). Figure 4.Diagnosis. – Moranopteris rupicola similis Moranopteridi

taenifoliae sed a qua habitu rupiculous et pilis ramosis cum 1 vel 2 ramis (vs. habitu epiphyticus, pilis non-ramosis et pilis ramosis cum 1 vel 2 ramis setaceis) differt.

Description. – Plants epipetric. Rhizomes erect with api-cal scales, the scales 2.1–2.7 × 0.3–0.4 mm, yellow-brown, lanceolate, bases slightly cordate, apices acute, mostly with apical or subapical gland-like cells, margins with long seti-form projections, or with also small lateral projections, rarely setiform projections occurring on the scale surfaces, setiform projections red-brown, 0.15–0.30 mm long. Leaves erect to arcuate, determinate, 5.5–13.0 × 0.45–1.00 cm, sparsely setose; stipes 0.5–1.0 cm long, yellow-brown to dark-brown, narrowly

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Fig. 4. Moranopteris rupicola. A, Habit; B, detail of the fertile segments showing sori and setae abaxially; C, detail of the segments showing setae and hydathodes adaxially; D, detail of the venation (cleared leaf); E, rhizome scale; F, detail of the apex of the rhizome scale; G, unbranched and branched hairs from the costa (re-hydrated hairs). A–C, E, F: Liesner 23289 (MO); D, G: Steyermark & al. 115650 (UC).

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Fig. 5. Moranopteris simplex. A, Habit; B, detail of the fertile segments showing sori and setae abaxially; C, detail of the segments showing setae and hydathodes adaxially; D, detail of the venation (cleared leaf); E, rhizome scale; F, unbranched and branched hairs from the costa (re-hydrated hairs). A–C, E: Mägdefrau 464 (B); D, F: Quijada 4 (VEN).

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Akaike, H. 1973. Information theory as an extension of the maximum likelihood principle. Pp. 267–281 in: Petrov, B.N. & Csaki, F. (eds.), Second International Symposium on Information Theory. Buda-pest: Akademiai Kiado.

winged; laminae linear, pinnatisect, gradually tapering proxi-mally to a narrow wing, chartaceous; costae visible or slightly visible with dark sclerenchyma abaxially, slightly visible with dark sclerenchyma adaxially; segments slightly ascending, 60°–80° to costae, 17–36 pairs, spaced (0.4–)0.6–1.8 mm, sub-opposite, gibbous, asymmetric, 2.8–4.9 × 1.9–2.9 mm, oblong, decurrent at bases, apices rounded to slightly acute, margins plane, without hyaline cells, or inconspicuous; veins usually furcate, at least in fertile segments, or sometimes simple in ster-ile segments, obscure, (1)2 hydathodes per segment, obovate or oblong; setae 1.6–2.4 mm long, dark red-brown, on both sides of the stipes, costae, and laminar tissue; hairs present on the costae and laminar tissue, rarely on the margins (mainly near to costae abaxially), hyaline or slightly red-brown, branched or rarely unbranched, the branched hairs mostly 1- or 2-branched, the main branch 2- or 3-celled, 0.15–0.25 mm long, 1st branch 1- or 2-celled, or sometimes 2-celled in the same insertion point, (<)0.10–0.15 mm long, 2nd branch, if present, 1-celled, < 0.1 mm long, the unbranched portion, 1-celled, < 0.1 mm long. Sori one per segment, rounded, 1.5–1.8 mm diam.

Additional specimens examined. – VENEZUELA. Bolívar, Cumbre del Ptari-tepuí, al norte de la Misión de Santa Teresita de Kavanayén, 2360–2420 m, 05°45′ N, 61°45′ W, 23 Feb 1978, Steyermark & al. 115650 (GH, UC, VEN).

Distribution. – Guyana and Venezuela, to be expected in northern Brazil; 2360–2800 m.

Discussion. – Moranopteris rupicola has rhizome scales with red-brown, long, setiform projections on the margins, sparsely setose leaves, gibbous segments, furcate veins, usu-ally two hydathodes per segment, and mostly branched hairs mainly near and on the costae (Fig. 4). Moranopteris rupicola resembles M. taenifolia by the habit and by the position of the hairs mostly occurring near and on the costae abaxially. However, M. taenifolia differs in having unbranched hairs and branched hairs with a setiform branch.

Etymology. – The specific epithet refers to the rocky habi-tat of the species in the Venezuelan tepuis.

2. Moranopteris simplex R.Y. Hirai & J. Prado, sp. nov. – Type: VENEZUELA. Aragua: Maracay, Choroní-Passes, 1600 m, 28 Feb 1958, K. Mägdefrau 464 (holotype: B-200077185). Figure 5.Diagnosis. – Species quam maxime affinis Moranopteridi

trichomanoidea, sed plerumque pilis non-ramosis et pilis furca-tis sparsis in lamina abaxialiter (vs. pilis ramosis cum 2–4-ra-mis, approximatis et in costa abaxialiter) differt.

Description. – Plants epiphytic. Rhizomes erect with api-cal scales, the scales 2.0–2.2 × 0.20–0.25 mm, yellow-brown, lanceolate to linear-lanceolate, bases slightly cordate, apices acute, with apical or subapical gland-like cells, margins en-tire or with small lateral projections, simple or furcate. Leaves erect to arcuate, determinate, 3.5–7.6 × 0.50–0.75 cm, moder-ately setose; stipes almost absent, brown to dark-brown, nar-rowly winged; laminae linear, pinnatisect, sometimes deeply pinnatifid, gradually tapering proximally to a narrow wing, chartaceous; costae obscured by laminar tissue abaxially and adaxially, sometimes slightly visible with dark sclerenchyma

abaxially; segments slightly ascending, ca. 60°–80° to costae, 18–35 pairs, spaced ca. 1.1–1.7 mm, subopposite, gibbous, asymmetric, 2.5–3.8 × 1.1–1.6 mm, oblong to oblong-lanceo-late, deltate toward bases, decurrent at base, apices rounded to slightly acute, margins slightly revolute, without hyaline cells; veins usually furcate, at least in fertile segments, or sometimes simple in sterile segments, obscure, (1)2 hydathodes per seg-ment, rarely 3 hydathodes, elliptic or obovate, sometimes round; setae 1.1–2.7 mm long, dark red-brown, on both surfaces of the costae and laminar tissue; hairs scattered all along the costae, laminar tissue abaxially, and margins, hyaline to yellowish or slightly red-brown, unbranched and branched, the unbranched hairs 2-celled, 0.05–0.15 mm long, the branched hairs 1- or 2-branched, mostly 1-branched, the main branch 2- or 3-celled, (<)0.1(–0.2) mm long, 1st branch 1- or 2-celled, 0.10–0.15 mm long, 2nd branch, if present, 1-celled, (<)0.1 mm long. Sori one per segment, round or nearly so, 1.4–1.8 mm diam.

Additional specimens examined. – VENEZUELA. Ara-gua, Colonia Tovar, s.d., Moritz s.n. p.p. (P-637667, P-696244); Distrito Federal, Caracas, Monte Grappa, 18 Aug 1985, Quijada 4 (VEN); El Junquito, 1925 m, 26 Mar 1943, Killip & Rohl 37170 (US).

Distribution. – Endemic to Venezuela; 1600–1925 m.Discussion. – Moranopteris simplex is easily distinguished

by hairs that are mostly yellowish, unbranched and 1-branched, scattered all along on the laminar tissue abaxially (Fig. 5).

The most similar species is Moranopteris trichomanoides, which occurs in the Greater Antilles and Central America. That differs by having mostly 2–4-branched hairs, near and on the costae abaxially, unbranched hairs absent.

Etymology – The specific epithet refers to the hairs which are less branched when compared to those of the most similar species (Moranopteris trichomanoides).

ACKNOWLEDGMENTS

This study was largely funded by a grant to the first author from Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, grant no. 06/06215-5). We thank The New York Botanical Garden for providing herbarium and other facilities during the visit of the first author, Judith Garrison Hanks, and Luciana Benjamim Bena-tti for SEM images (setae and spores, respectively), and Alejandra Vasco, Luz A. Triana-Moreno, Michael Kessler, Marcus Lehnert, and Michael Sundue for providing us with silica-dried material. Finally, we are especially grateful to Dr. Robbin Moran, whose support was fundamental to the development of this study (grant from the United States National Science Foundation, DEB 0717056). We also thank the two anonymous reviewers for their comments on the manuscript.

LITERATURE CITED

1136

TAXON 60 (4) • August 2011: 1123–1137Hirai & al. • New Neotropical fern genus Moranopteris

Desalle, R., Giribert, G. & Wheeler, W. 2002. Techniques in molecu-lar systematics and evolution. Basel: Birkhäuser Verlag.

Edgar, R.C. 2004. Muscle: Multiple sequence alignment with high accuracy and high throughput. Nucl. Acids Res. 32: 1792–1797.

Farrar, D.R. 1967. Gametophytes of four tropical fern genera repro-ducing independently of their sporophytes in the southern Ap-palachians. Science 155: 1266–1267.

Felsenstein, J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution 39: 783–791.

Hasebe, M., Omori, T., Nakazawa, M., Sano, T., Kato, M. & Iwat-suki, K. 1994. rbcL gene sequences provide evidence for the evo-lutionary lineages of leptosporangiate ferns. Proc. Natl. Acad. Sci. U.S.A. 91: 5730–5734.

Hasebe, 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.H., Haufler, C.H. & Hauk, W.D. 1995. Fern phylogeny based on rbcL nucleotide sequences. Amer. Fern J. 85: 134–181.

Huelsenbeck, J.P. & Ronquist, F. 2001. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754–755.

Kraus, J.E. & Arduin, M. 1997. Manual básico de métodos em mor-fologia vegetal. Seropédica, Rio de Janeiro: Edur.

Labiak, P.H. & Matos, F.B. 2007. A new hybrid and two new com-binations in neotropical grammitid ferns. Brittonia 59: 182–185.

Labiak, P.H., Rouhan, G. & Sundue, M. 2010a. Phylogeny and tax-onomy of Leucotrichum (Polypodiaceae): A new genus of gram-mitid ferns from the Neotropics. Taxon 59: 911–921.

Labiak, P.H., Sundue, M. & Rouhan, G. 2010b. Molecular phylogeny, character evolution, and biogeography of the grammitid fern genus Lellingeria (Polypodiaceae). Amer. J. Bot. 97: 1354–1364.

Lehnert, M., Kessler, M., Schmidt-Lebuhn, A.N., Klimas, S.A., Fehlberg, S.D. & Ranker, T.A. 2009. Phylogeny of the fern ge-nus Melpomene (Polypodiaceae) inferred from morphology and chloroplast DNA analysis. Syst. Bot. 34: 17–27.

Meier, R., Kwong, S., Vaidya, G. & Ng, P.K.L. 2006. DNA barcoding and taxonomy in Diptera: A tale of high intraspecific variability and low identification success. Syst. Biol. 55: 715–728.

Mickel, J.T. & Smith, A.R. 2004. The pteridophytes of Mexico. Mem. New York Bot. Gard. 88: 1–1054.

Parris, B.S. 1990. Grammitidaceae. Pp. 153–157 in: Kramer, K.U. & Green, P.S. (eds.), The families and genera of vascular plants, vol. 1, Pteridophytes and gymnosperms. Berlin: Springer.

Parris, B.S. 1998. Chrysogrammitis, a new genus of Grammitidaceae (Filicales). Kew Bull. 53: 909–918.

Parris, B.S. 2009. New genera of Malesian Grammitidaceae (Mo-nilophyta). Blumea 54: 217–219.

Philippe, H. 1993. MUST, a computer package of management utilities for sequences and trees. Nucl. Acids Res. 21: 5264–5272.

Posada, D. 2008. jModelTest: Phylogenetic model averaging. Molec. Biol. Evol. 25: 1253–1256.

Proctor, G.R. 1985. Ferns of Jamaica. London: British Museum of Natural History.

Pryer, K.M., Smith, A.R. & Skog, J.E. 1995. Phylogenetic relation-ships of extant ferns based on evidence from morphology and rbcL sequences. Amer. Fern J. 85: 205–282.

Rambaut, A. & Drummond, A.J. 2004. Tracer. Oxford: University of Oxford.

Ranker, T.A., Geiger, J.M., Kennedy, S.C., Smith, A.R., Haufler, C.H. & Parris, B.S. 2003. Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae). Molec. Phylog. Evol. 26: 337–347.

Ranker, T.A., Smith, A.R., Parris, B.S., Geiger, J.M.O., Haufler, C.H., Straub, S.C.K. & Schneider, H. 2004. Phylogeny and evo-lution of grammitid ferns (Grammitidaceae): A case of rampant morphological homoplasy. Taxon 53: 415–428.

Ranker, T.A., Sundue, M., Labiak, P.H., Parris, B. & Rouhan, G. 2010. New insights into the phylogeny and historical biogeogra-phy of the Lellingeria myosuroides clade (Polypodiaceae). Version 48. PLoS Currents: Tree of Life. 18 Nov. 2010 [revised 22 Nov. 2010]: PMC2989831. http://knol.google.com/k/tom-a-ranker/new -insights-into-the-phylogeny-and/uggrdrq3hh35/4?collectionId=28qm4w0q65e4w.46# (accessed 29 Nov 2010).

Ronquist, F. & Huelsenbeck, J.P. 2003. MrBayes 3: Bayesian phyloge-netic inference under mixed models. Bioinformatics 19: 1572–1574.

Schneider, H., Smith, A.R., Cranfill, R., Hildebrand, T.J., Haufler, C.H. & Ranker, T.A. 2004. Unraveling the phylogeny of poly-grammoid ferns (Polypodiaceae and Grammitidaceae): Exploring aspects of the diversification of epiphytic plants. Molec. Phylog. Evol. 31: 1041–1063.

Schuettpelz, E. & Pryer, K.M. 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037–1050.

Smith, A.R. 1992. A review of the fern genus Micropolypodium (Gram-mitidaceae). Novon 2: 419–425.

Smith, A.R. 1993. Terpsichore, a new genus of Grammitidaceae (Pte-ridophyta). Novon 3: 478–489.

Smith, A.R. 1995. Grammitidaceae. Pp. 366–367 in: Moran, R.C. & Riba, R. (eds.), Psilotaceae a Salviniaceae. in: Davidse, G., Sousa, M.S. & Knapp, S. (eds.), Flora Mesoamericana. Ciudad de México: Universidad Nacional Autónoma de México.

Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A classification for extant ferns. Taxon 55: 705–731.

Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. Pp. 417–467 in: Ranker, T.A. & Haufler, C.H. (eds.), Biology and evolution of ferns and lycophytes. Cambridge: Cambridge University Press.

Sundue, M.A., Islam, M.B. & Ranker, T.A. 2010. Systematics of grammitid ferns (Polypodiaceae): Using morphology and plastid sequence data to resolve the circumscription of Melpomene and the polyphyletic genera Lellingeria and Terpsichore. Syst. Bot. 35: 701–715.

Swofford, D.L. 2002. PAUP*: Phylogenetic analyses using parsimony (*and other methods), version 4. Sunderland, Massachusetts: Sinauer.

Taberlet, P., Gielly, L., Pautou, G. & Bouvert, J. 1991. Universal primers for amplification of three non-coding regions of chloro-plast DNA. Pl. Molec. Biol. 17: 1105–1109.

Tryon, A.F. & Lugardon, B. 1990. Spores of the Pteridophyta. New York: Springer.

Appendix. Species, vouchers (collector, collector number, and herbarium), locality, and GenBank accession numbers of the DNA sequences used in this paper (rbcL, atpB, trnL-F). Missing data: –. Asterisks indicate newly obtained sequences.

Adenophorus tripinnatifidus Gaudich., Ranker 1102 (COLO), Hawaiian Islands, U.S.A., AF468207, AF469783, AF469796; Ascogrammitis pichinchae (Sodiro) Sundue, Dassler 94-7-13-1 (ILLS), Colombia, AY460675, AY459508, GU476711; Calymmodon gracilis (Fée) Copel., Chiou 97-09-12-01 (COLO, TAIF, UC), Taiwan, AY362341, AY459451, GU476618; Ceradenia spixiana (Mart. ex Mett.) L.E. Bishop, Salino 3008 (UC), Brazil, AY460623, AY459457, –; Chrysogram-mitis glandulosa (J. Sm.) Parris, Ranker 2195 (BORH, SAN, SNP, COLO), Malaysia, JF514014*, JF514082*, JF514048*; Chrysogrammitis musgraviana (Baker) Parris, Kessler 12570 (UC), Sabah, Malaysia, AY460624, AY459458, GU476630; Cochlidium serrulatum (Sw.) L.E. Bishop, Hirai & Schwartsburd 541 (SP), Brazil, JF514010*, JF514078*, JF514044*; Cochlidium punctatum (Raddi) L.E. Bishop, Silva 3914 (UC), Brazil, JF513987*, JF514057*, GU476631; Ctenopterella denticulata (Blume) Parris, Ranker 2113 (BORH, SAN, SNP, COLO), Malaysia, JF514013*, JF514081*, JF514047*; Ctenopteris heterophylla (Labill.) Tindale, Parris 12419 (AK), New Zealand, AY460629, AY459462, –; Ctenopteris lasiostipes (Mett.) Brownlie, Hodel 1448 (UC), New Caledonia, AY460630, AY459463, –; Dasygrammitis crassifrons (Baker) Parris, Game 95-80 (UC), Fiji, JF513992*, JF514062*, JF514026*; Enterosora percrassa (Baker) L.E. Bishop, Moraga

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& Rojas 508 (UC), Costa Rica, AY460635, AY459468, GU476636; Grammitis billardierei Willd., Parris 12421 (AK), New Zealand, AY460637, AY459469, JF514042*; Grammitis bryophila (Maxon) F. Seym., Rojas & al. 3240 (UC), Costa Rica, AF468208, AF469784, AF469797; Grammitis deplanchei (Baker) Copel., Hodel 1450 (UC), New Caledonia, AY460639, AY459471, –; Grammitis poeppigiana (Mett.) Pic.Serm., Weber 13772 (COLO), Australia, AY460647, Taylor 6072 (UC), Chile, AY459479, –; Grammitis tenella Kaulf., Ranker 1352 (COLO); Hawaiian Islands, U.S.A., AF468198, AF469773, AF469786; Lellin-geria apiculata (Kunze ex Klotzsch) A.R. Sm. & R.C. Moran, Salino 3009 (UC), Brazil, AY362343, AY459480, GU476642; Lellingeria limula (Christ) A.R. Sm. & R.C. Moran, Sundue 1736 (INB, NY, UC, UPCB), Costa Rica, GU476903, GU476765, GU476651; Leucotrichum schenckii (Hieron.) Labiak, Salino 4547 = 4538 (UC, BHCB), Brazil, AY460651, AY459483, GU476657; Melpomene flabelliformis (Poir.) A.R. Sm. & R.C. Moran, Sanchez 183 (UC), Colombia, AY460656, AY459488, –; Melpomene moniliformis (Lag. ex Sw.) A.R. Sm. & R.C. Moran, Moraga & Rojas 446 (INB), Costa Rica, AY460654, AY459486, GU476664; Microgramma bifrons (Hook.) Lellinger, Neill & al. 8309 (UC), Ecuador, AY362582, van der Werff 18062 (UC), Peru, EF463499, DQ642224; Microgramma percussa (Cav.) de la Sota, Smith 1357 (UC), Venezuela, AY362574, AY459516, GU476669; Micropolypodium okuboi (Yatabe) Hayata, Par-ris 12154 (AK), Japan, JF513994*, JF514064*, JF514028*; Kanebira & Sasaki 21672 (UC), Formosa, JF513998*, –, –; Wilson 2387 (UC), Japan, JF514018*, JF514086*, JF514053*; Micropolypodium sikkimense (Hieron.) X.C. Zhang, Miehe 00-093-32 (UC), Bhutan, JF513999*, JF514068*, JF514032*; Xian & al. s.n. (UC), China, JF514019*, JF514087*, JF514054*; Moranopteris achilleifolia (Kaulf.) R.Y. Hirai & J. Prado, Cordeiro & Ribas 1398 (MBM, UC), Brazil, AY460666, AY459499, –; Moranopteris aphelolepis (C.V. Morton) R.Y. Hirai & J. Prado, Borke & Jaramillo 2701 (NY), Ecuador, –, –, JF514021*; Jiménez & Vidarre 557 (UC), Bolivia, JF513996*, JF514066*, JF514030*; Moranopteris basiattenuata (Jenman) R.Y. Hirai & J. Prado, Breedlove & Thorne 30160 (NY), Mexico, JF513988*, JF514058*, JF514022*; Moranopteris blepharidea (Copel.) R.Y. Hirai & J. Prado, Jiménez 708 (GOET), Bolivia, JF513995*, JF514065*, JF514029*; Moranopteris caucana (Hieron.) R.Y. Hirai & J. Prado, van der Werff & al. 13445 (UC), Ecuador, JF513991*, JF514061*, JF514024*; Lehnert 182 (GOET), Ecuador, JF514002*, JF514071*, JF514035*; Moranopteris cookii (Underw. & Maxon) R.Y. Hirai & J. Prado, Sundue & al. 1771 (NY), Costa Rica, JF514007*, JF514076*, JF514040*; Moranopteris gradata (Baker) R.Y. Hirai & J. Prado, Hirai & al. 537 (SP), Brazil, JF514009*, JF514077*, JF514043*; Mo-ranopteris grisebachii (Underw. ex C. Chr.) R.Y. Hirai & J. Prado, Maxon 9961 (NY), Jamaica, JF514008*, –, JF514041*; Moranopteris hyalina (Maxon) R.Y. Hirai & J. Prado, Rojas & al. 3210 (CR, INB, MO, UC), Costa Rica, AY362344, AY459490, GU476670; Lehnert 1426 (GOET), Ecuador, JF514001*, JF514070*, JF514034*; Sundue 1148 (NY), Ecuador, JF514006*, JF514075*, JF514039*; Moranopteris longisetosa (Hook.) R.Y. Hirai & J. Prado, Rojas & al. 3209 (CR, INB, MO, UC), Costa Rica, AY460674, AY459507, –; Lehnert 596 (GOET), Bolivia, JF514003*, JF514072*, JF514036*; Moranopteris microlepis (Rosenst.) R.Y. Hirai & J. Prado, Smith 2584 (UC), Costa Rica, JF513986*, JF514056*, JF514020*; Kluge 2003 (UC), Costa Rica, JF513997*, JF514067*, JF514031*; Moranopteris nana (Fée) R.Y. Hirai & J. Prado, Diaz & al. 4747 (NY), Venezuela, JF513990*, –, –; Moranopteris perpusilla (Maxon) R.Y. Hirai & J. Prado, Hirai & al. 574 (SP), Brazil, JF514011*, JF514079*, JF514045*; Moranopteris plicata (A.R. Sm.) R.Y. Hirai & J. Prado, Lehnert 929 (GOET, UC), Ecuador, JF514005*, JF514074*, JF514038*; Moranopteris serricula (Fée) R.Y. Hirai & J. Prado, Feldmann s.n. (P), Guadeloupe, –, –, JF514025*; Wilbur & al. 8084 (NY), Dominica, JF514017*, JF514085*, JF514052*; Moranopteris setosa (Kaulf.) R.Y. Hirai & J. Prado, Hirai & al. 599 (SP), Brazil, JF514012*, JF514080*, JF514046*; Moranopteris taenifolia (Jenman) R.Y. Hirai & J. Prado, Rothfels 08-116 (DUKE), Costa Rica, –, JF514060*, JF514023*; Triana-Moreno s.n. (SP), Costa Rica, JF514015*, JF514083*, JF514050*; Moranopteris trichomanoides (Sw.) R.Y. Hirai & J. Prado, Gomez 114 (NY), Honduras, JF513993*, JF514063*, JF514027*; Zanoni & al. 45973 (NY), Dominica, –, –, JF514055*; Moranopteris truncicola (Klotzsch) R.Y. Hirai & J. Prado, Lehnert 862 (GOET), Ecuador, JF514004*, JF514073*, JF514037*; Vasco & Sundue 626 (NY), Colombia, JF514016*, JF514084*, JF514051*; Moranopteris williamsii (Maxon) R.Y. Hirai & J. Prado, Kessler & al. 7173 (UC), Bolivia, JF514000*, JF514069*, JF514033*; Moranopteris zurquina (Copel.) R.Y. Hirai & J. Prado, Rojas & Mata 3021 (UC), Costa Rica, AY460659, AY459492, –; Oreogrammitis hookeri (Brack.) Parris, Ranker 1116 (COLO), Hawaiian Islands, U.S.A., AY460642, AY459473, EF178655; Pecluma eurybasis (C. Chr.) M.G. Price, Danton s.n. (GOET), Bolivia, EF463255, EF463504, Kessler s.n. (GOET), Bolivia, FJ825691; Polypodium vulgare L., Schneider s.n. (GOET), Germany, EF551065, EF463510, EF551119; Prosaptia contigua (G. Forst.) C. Presl, Chiou 97-09-12-05 (TAIF, COLO, UC), Taiwan, AY362345, AY459494, EF178663; Prosaptia nutans (Blume) Mett., Ranker & Trapp 1765 (COLO, UC), Papua New Guinea, AY460631, AY459464, –; Radiogrammitis parva (Brause) Parris, Ranker 1763a (COLO, UC), Papua New Guinea, AY460644, AY459476, –; Scleroglossum sulcatum (Kuhn) Al-derw., Bowden-Kerby in Raulerson 24182b (GUAM, UC), Pohnpei, AY460665, AY459498, JF514049*; Serpocaulon triseriale (Sw.) A.R. Sm., Jiménez 1994 (UC), Bolivia, DQ151926, EF463516, DQ151980; Terpsichore asplenifolia (L.) A.R. Sm., Moraga & Rojas 506 (INB), Costa Rica, JF513989*, JF514059*, –; Terpsichore cultrata (Willd.) A.R. Sm., Dassler 94-7-19-1 (ILLS), Colombia, AY460669, AY459502, –; Terpsichore lanigera (Desv.) A.R. Sm., Léon 3647 (USM, UC), Peru, AY460672, AY459505, GU476718; Terpsichore lehmanniana (Hieron.) A.R. Sm., Wilson 2589 (UC), Ecuador, AY460673, AY459506, –; Themelium conjunctisorum (Baker) Parris, Ranker & Trapp 1758 (COLO, UC), Papua New Guinea, AY460680, AY459514, –; Tomophyllum repandulum (Mett.) Parris, Ranker & Trapp 1767 (COLO, UC), Papua New Guinea, AY460633, AY459466, –.

Appendix. Continued.