Corresponding author: Thaís Elias Almeida ( email@example.com )
Academic editor: Blanca Leon
© 2017 Thaís Elias Almeida, Alexandre Salino, Jean-Yves Dubuisson, Sabine Hennerquin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Almeida TE, Salino A, Dubuisson J-Y, Hennequin S (2017) Adetogramma (Polypodiaceae), a new monotypic fern genus segregated from Polypodium. PhytoKeys 78: 109-131. https://doi.org/10.3897/phytokeys.78.12189
Polypodiaceae is one of the most diverse and abundant families of ferns in tropical and subtropical forests. Despite multiple studies investigating its phylogeny and taxonomy, several generic boundaries within the family still need clarification. One of the most problematic circumscriptions is that of Polypodium L., and one species that still contributes to this uncertainty is Polypodium chrysolepis Hook. The main goal of this study was to use molecular and morphological data to clarify the relationships of P. chrysolepis inside the polygrammoid clade. Sequences from three plastid regions (cpDNA – rbcL, rps4 and rps4–trnS IGS) from fifty species belonging to thirty-two genera of Polypodiaceae were analyzed using maximum likelihood and Bayesian inference. Polypodium chrysolepis constitutes an isolated lineage among the neotropical polygrammoid ferns, close to Serpocaulon and the grammitids, and is recognized here in a new genus. It can be distinguished by its entire leaves with free veins and peltate, pedicellate, lanceolate paraphyses. A new combination, Adetogramma chrysolepis, is proposed and a new taxonomic treatment is presented; its conservation status was assessed using IUCN Red List Categories and Criteria.
Andes, Polypodiaceae, phylogeny, Serpocaulon, taxonomy
Polypodiaceae is one of the richest fern families, and one of the most diverse and abundant groups of vascular plants in tropical and subtropical forests (
One of the most problematic circumscriptions is that of Polypodium L. (
One species that still contributes to this uncertainty is Polypodium chrysolepis Hook., a species occurring in the Andes from northern Argentina to Ecuador. The generic placement of this species has been controversial: it was described in Polypodium by
The main goal of this study was to employ molecular and morphological data to investigate the relationships of Polypodium chrysolepis within the polygrammoid clade, and to use available morphological and phylogenetic information to formally propose an adequate generic placement for this species in Polypodiaceae.
Fifty species from thirty-two genera (sensu
Total DNA was extracted from field-acquired silica gel-dried or fresh tissues, using the Qiagen DNeasy Plant mini kit (Qiagen Inc., Valencia, CA, USA). PCR amplifications were performed for two chloroplast genome regions: rbcL (coding region; ca. 1,300 bp) and rps4 (the coding region rps4 and the intergenic spacer rps4–trnS; ca. 1,100 bp). Amplifications were done in a single reaction with primers 1F and 1365R for the rbcL region (
Polymerase chain reactions were performed in a 20 μL solution containing 1.0 μL of non-diluted genomic DNA template, 2.0 μL of PCR buffer (Qiagen 10 × PCR Buffer), 1.0 μL of DMSO, 1.0 μL of BSA (4 mg/mL), 0.8 μL of dNTPS (10 mM), 0.32 μL (10 μM) of each primer, 0.12 units of Taq Dna polymerase (Qiagen, 5 units μL), and 14.44 μL of ultra-pure water. Thermal cycling conditions were the same for both regions: 3 min at 94°C, 35 cycles of 45 s at 94°C, 60 s at 53°C and 90 s at 72°C, followed by 5 min at 72°C. Amplicons were purified by precipitation with polyethylene glycol (PEG) and sequenced by Macrogen (Seoul, South Korea) in a bidirectional sequencing reaction in an ABI3730XL.
Sequence electropherograms were edited using the STADEN package software (
Datasets were analyzed using maximum likelihood (ML) and Bayesian inference (BI). Maximum likelihood was performed using IQ-TREE (
Selected models and parameter values for data partitions used in this study.
|Substitution model (rate matrix)|
Taxonomic conclusions were based on the study of specimens from the following herbaria: BHCB, BM, BR, G, GH, K, LPB, M, MO, NY, P, Q, QCA, QCNE, QPLS, US, USM, and USZ. Specimens with barcode are cited with herbarium acronym followed by barcode number. Abbreviation of genera and species followed IPNI (ipni.org) and morphological terms follow
The final combined dataset presented 2339 bp, 599 from rps4 gene, 473 from the rps4-trnS IGS, and 1267 from rbcL. All analyses recovered the main polygrammoid clades found by
Inside the neotropical clade, Synammia appears as sister to all the other neotropical clades, a result also obtained by
Majority rule consensus tree resultant from Bayesian inference analyses from combined rbcL and rps4 datasets. Values above branches nodes correspond to Bayesian posterior probabilities and maximum likelihood bootstrap support, respectively. +/+ indicates with 1.00 PP and 100% BS support. Clades cited in text follow
The polygrammoid topology recovered in our analyses agrees with previous results from several studies (
In our analyses, Serpocaulon, a group segregated from Polypodium (
Comparison of character states among Adetogramma and the morphologically or phylogenetically closest genera/groups.
|Adetogramma||Serpocaulon 1||Grammitids||Pecluma||Polypodium 2||Pleopeltis 3||Microgramma 4|
|Rhizome||Long-creeping, branched||Long- to short-creeping, sparingly branched||Short-creeping to erect, usually unbranched||Long- to short-creeping, unbranched||Long- to short-creeping, branched||Long- to short-creeping, branched||Long-creeping, branched|
|Rhizome scales||Peltate, non-clathrate||Peltate, clathrate||Basifixed, non-clathrate or clathrate, or absent||Basifixed to peltate, non-clathrate||Peltate, non-clathrate||Peltate, non-clathrate, to clathrate at margins||Peltate, non-clathrate|
|Fronds||Monomorphic||Monomorphic||Monomorphic, or the distal fertile portion modified||Monomorphic||Monomorphic||Monomorphic to dimorphic||Monomorphic to dimorphic|
|Lamina||Simple||Pinnatifid to pinnate, rarely simple||Simple to 3-pinnate||Pinnatisect to pinnate||Deeply pinnatifid to pinnate||Simple to pinnatifid, rarely pinnate-pinnatifid or more divided||Simple to lobate|
|Indument on lamina||Scales||Glabrous, trichomes, or scales (confined to costae and rachises)||Trichomes, and sometimes glands||Trichomes||Glabrous or with trichomes||Scales||Glabrous, trichomes
|Veins||Free||Regularly anastomosing (goniophlebioid), areoles with one included veinlet||Usually free, sometimes anastomosing with or without included free veinlets||Free, rarely anastomosing, but never reticulate||Free to anastomosing, with one single included veinlet||Free to anastomosing, areoles with 1-3 free or netted included veins||Anastomosing, with simple included veinlets|
|Sori||Round to oblong, 1 row between costa and margins||Round, 1-10 rows between costa and margins||Round to elongate, 1 row between costa and margins, or confluent||Round to oblong, 1 row between costa and margins||Round to oblong, 1-5 rows between costa and margins||Round to oblong, or linear, rarely marginal and coalescing, in 1 row between costa and margins||Round to elongate, 1 row between costa and margins, confluent or forming several irregular rows between costa and margins|
|Paraphyses||Peltate, pedicellate, scales||Absent or short 2-3-celled glands||Present or absent||Present, glandular trichomes||Absent or if present, filamentous or branched||Absent, or round peltate,||Trichomes or sessile scales, or absent|
|Spores||Bilateral, monolete, verrucate||Bilateral, monolete, verrucate, generally tuberculate, occasionally winged||Tetrahedral-globose, trilete||Bilateral,
monolete, smooth to tuberculate
|Bilateral, monolete||Bilateral, monolete, shallowly to prominently verrucate||Bilateral, monolete, tuberculate|
Adetogramma chrysolepis (Hook.) T.E.Almeida. A Habit B Rhizome scale C Sterile lamina detail showing free venation and laminar scales D Fertile lamina detail showing sori, paraphyses, and laminar scales E Paraphysis detail. Drawings A–D by H. Fukuda from Dorr et al. 6764 (NY); drawing E by Juliana Ventura from T.E. Almeida & L.L. Giacomin 3121 (BHCB). Scale bars: A = 5 cm, B–E = 1 mm.
Polypodium chrysolepis was combined by
Grammitid ferns, the lineage sister to the Polypodium chrysolepis+Serpocaulon clade, are a very distinct group of species inside the polygrammoid ferns. Once recognized as a separated family (
Polypodium s.s., following
Our results do not support the inclusion of Polypodium chrysolepis in any genus previously recognized, including Microgramma, Pleopeltis, or Polypodium s.s. Therefore, we consider this species as constituting a separated, isolated lineage inside the polygrammoid clade. Because the species also has a morphology distinct from that of all other known genera in Polypodiaceae, we believe it merits recognition as a genus, and is described below.
Adetogramma is similar to Microgramma and Pleopeltis in its epiphytic habit, long-creeping rhizomes and in having entire leaves with one row of sori on each side of the midrib, but differs from these genera by having free veins (vs. veins anastomosing in Microgramma and Pleopeltis) and peltate, pedicellate, lanceolate paraphyses (vs. hairlike or lanceolate and sessile paraphyses in Microgramma, and round, peltate, pedicellate paraphyses in Pleopeltis).
Adetogramma chrysolepis (Hook.) T.E.Almeida, comb. nov., Polypodium chrysolepis Hook., Icon. Pl. 8: t. 721. 1845.
Lepicystis chrysolepis (Hook.) Diels, Nat. Pflanzenfam. 1(4): 322, f. 167A–B. 1899.
Type: Based on Polypodium chrysolepis Hook.
Microgramma chrysolepis (Hook.) Crabbe, Brit. Fern Gaz. 9: 316. 1967.
Type: Based on Polypodium chrysolepis Hook.
Polypodium bangii Baker, Bull. Misc. Inform. Kew 1901: 145. 1901.
Type: Bolivia. Yungas, 1890, A.M. Bang 734 (lectotype, designated here: BM! [BM000936895]; isolectotypes: B! [B200075587], BR!, GH!, K! [K000590773], LE!, MO! [MO5472871)], NY! [NY00144786, NY00144787], US! [US00065725]).
Polypodium chrysolepis Hook., Icon. Pl. 8: t. 721. 1845.
Ecuador. Andes de Quito, W. Jameson 37 (wrongly typed in protologue as “73”; lectotype, designated by
Plants epiphytic or epipetric, rarely terrestrial. Rhizomes long-creeping, branched, 0.6–0.9 mm wide, cylindrical, with four vascular bundles; short, perpendicular roots about 5–20 mm long, these regularly spaced, covered with brownish root hairs; rhizome scales 4.3–7.1 mm long, peltate, not clathrate, linear-lanceolate, with elongate cells, the margins entire from the base to the middle and toothed beyond the middle, with 1- or 2-celled marginal teeth, scales concolorous, stramineous and usually darker at the attachment point. Fronds remote, 2.2–4.5 mm apart, articulate, monomorphic. Stipes nearly absent to 55 mm long, 0.4–0.7 mm in diameter, covered with sparse peltate, lanceolate, sessile, non-clathrate, concolorous, stramineous scales, these 1.4–3.5 mm long, darker at the attachment point; phyllopodia darker than stipes. Laminae light green, 5.0–17.0 × 1.0–2.5 cm, simple, chartaceous, linear-lanceate to lanceolate, bases acuminate to attenuate, decurrent in the distal third of the stipe, apices acute to obtuse, laminar surfaces squamose on both sides, scales lanceolate, peltate, sessile, non-clathrate, concolorous, translucent, stramineous, darker at attachment point, slightly erose at bases and entire at apices, scales present also on abaxial and adaxial sides of costae and veins, and on the laminar margins, 1.3–2.6 mm long. Veins free, immersed, obscure, 1–2 furcate, not reaching laminar margins, midribs and lateral veins immersed on both sides of the laminae, not evident. Sori superficial, rounded to oblong, 1.6–2.6 × 1.9–5.0 mm, terminal to subterminal on veins, receptacles elongate, sporangia long-stalked, paraphyses present, scale-like, similar those of the laminar surfaces, peltate, pedicellate, with pedicels as long as those of the sporangia, paraphyses completely covering immature sori. Spores yellow, with verrucate surfaces.
Restricted to central and southern Andes, with known collections from Ecuador, Peru, Bolivia, and Argentina (Fig.
Least Concern (LC -
The generic epithet refers to the most distinctive character of the species, the free venation (Fig.
Adetogramma is a monotypic genus, and although its sole species, A. chrysolepis, varies in laminar size and shape and stipe length, all other characters, such as the rhizome and stipe scales, venation, and paraphyses, are constant. Specimens from the Argentinean Provinces of Tucumán and Salta and Bolivian Provinces of Tarija and Chuquisaca have very long, linear laminae, and longer stipes, but in other characters are congruent with the circumscription here adopted for A. chrysolepis. This variation may reflect colonization of drier, seasonal habitats in a subtropical region.
Morphologically, Adetogramma chrysolepis shares features with several neotropical genera of Polypodiaceae (Table
ARGENTINA. Jujuy: San Antonio, 27 Apr 2015, C. Martín 479 (SI); Valle Grande, 14 Apr 2016, C. Martín 730 (SI). Salta: Salta, 21 Nov 1945, S.A. Pieroffi 1326 (NY!); Santa Victoria, 05 Dec 2015, C. Martín 673 (SI). Tucumán: s.l., 1952, H. Brücher s.n. (LP!); s.l., s.d., L. Castillón 2248 (BM!); s.l., s.d., M. Lillo 11534 (BM!, GH!, K!, NY!); La Ventanita, s.d., M. Lillo 16713 (BM!, GH!, K!); Burruyacu, 20 Jan 1947, Borsini s.n. (LP!); Chicligasta, 11 Dec 1925, S. Venturi 4062 (BM!, GH!); idem, 13 Feb 1924, S. Venturi 3151 (GH!); Tafi del Vale, Jan 1912, L. Castillón 35 (BM!, GH!, K!, LP!, NY!). BOLIVIA. Chuquisaca: Hernando Siles, 09 Nov 2007, J. Villalobos 928 (MO!, UC). Cochabamba: Ayopaya, 07 May 1990, E. Hennipman 8148 (LPB!); idem, 28 Oct 2007, J. Terán 1486 (MO!); idem, 30 Oct 2007, J. Terán 1563 (MO!); Carrasco, 20 Mar 1991, I. Hensen 1819 (LPB!); idem, 25 Feb 1996, M. Mercado 501 (MO!); Chapare, 30 Sep 2001, J.R.I. Wood 17271 (LPB!); Cochabamba, 27 Jan 1950, W.M.A. Brooke 6081 (BM!, NY!); Jose Carrasco Torrico, 27 Jun 1996, M. Kessler 6763 (NY!, UC); idem, 02 Jul 1996, M. Kessler 6877 (NY!, UC); Tiraque, 10 May 2005, E. Zurita 390 (MO!). La Paz: Bautista Saavedra, 27 Apr 1993, P. Gutte 559 (LPB!); Franz Tamayo, 24 Feb 2008, A.F. Fuentes Claros 11982 (MO!, UC); idem, 04 Mar 1980, J. Krach 9207 (LPB!); idem, 06 Apr 2009, M.I.L. Rivera 599 (MO!); Inquisivi, 21 Dec 1989, L.J. Dorr 6764 (LPB!, NY!); idem, 18 Feb 1989, M.A. Lewis 35242 (LPB!, MO!, NY!, UC); idem, 09 Mar 1991, M.A. Lewis 38230 (MO!); idem, 04 Sep 1991, M.A. Lewis 39744 (F, GH!, LPB!, MO!); Larecaja, 1818, G. Mandon 1560 (BM!, G!, K!, NY!); Murillo, 16 May 1985, J.C. Solomon 13742 (LP!, LPB!, MO!, NY!); Nor Yungas, XI/1900, O. Buchtien 2751 (P!); idem, 07 Mar 1969, H. Doppelbaur s.n. (MO!); Pedro Domingo Murillo, 17 Mar 2012, T.E. Almeida 3121 (BHCB!, LPB!); Pongo, s.d., W.M.A. Brooke 5456 (BM!, NY!); Unduavi, 1890, A.M. Bang 734 (B!, BR!, GH!, K!, LE!, MO!, US!); idem, 30 Mar 1977, J.D. Boeke 1375 (NY!); idem, Feb 1914, O. Buchtien 420 (B!, BM!, G!, GH!, K!, NY!, P!); idem, 19 Jun 1912, E. Rosenstock 49 (B!, P!). Santa Cruz: Manuel M. Caballero, 11 Apr 2004, R. Nuñez Cabrera 682 (MO!, NY!, UC, USZ!); idem, 08 Mar 2012, T.E. Almeida 3083 (BHCB!, HSTM!, LPB!); s.l., 26 Oct 1928, J. Steinbach 8531 (BM!, GH!, K!, MO!, NY!). Tarija: Aniceto Arce Ruiz, 13 Jun 2004, I. Jimenez 2420 (NY!, UC); Arce, 10 Nov 2004, H. Huaylla 1509 (MO!); idem, 27 Jun 2005, H. Huaylla 1882 (MO!); idem, 12 Nov 2004, M. Serrano 5082 (MO!); idem, 12 Nov 2004, M. Serrano 5136 (MO!); idem, 03 Feb 2005, M. Serrano 5945 (MO!); O’Connor, 22 Oct 1983, S.G. Beck 9651 (LPB!); s.l, s.d., M. Cárdenas 1036 (GH!); s.l., 07 Mar 1969, H. Doppelbaur 291 (M!); s.l., 06 May 1928, [illegible]1569 (B!); s.l., 21 Aug 1926, [illegible] 2559 (B!). ECUADOR. Pichincha: Quito, s.d., W. Jameson 73 (G!, K!). PERU. Amazonas: s.l., 03 Aug 1962, J.J. Wurdack 1592 (GH!). Ancash: Carhuaz, 14 Feb 1985, D.N. Smith 9571 (MO!, USM); idem, 18 Jul 1985, D.N. Smith 11264 (BM!, ENAG, F, GH!, MICH, MO!, NY, UC, USM); Huari, 08 May 1986, D.N. Smith 12415 (BM!, F, LPB!, MO!, NY, UC); idem, 13 Jun 1986, Smith 12648 (MO!); Yungay, 13 Jan 1985, Smith 9150 (GH!, MO!, USM); idem, 17 Apr 1985, D.N. Smith 10326 (MO!, USM); idem, 19 Apr 1985, D.N. Smith 10463 (MO!, USM). Apurimac: Abancay region, Oct 1935, students of Santander C. s.n. (UC); Abancay, Tamburco, 6 Jun 2015, V. Zuñiga 452 (USM). Cusco: s.l., 31 May 2002, W.L. Galiano 4034 (MO!); s.l., 14 Feb 1987, P.Núñez Vargas 6988 (MO!); s.l., 10 Jun 2002, L.Valenzuela Gamarra 238 (NY!, MO!); La Convención, 11 Jul 2003, E. Bonino 832 (MO!); idem, 23 Jul 2003, E. Bonino 909 (MO!, UC); idem, 26 Mar 2004, I. Huamantupa 4428 (MO!); idem, 21 Sep 2005, I. Huamantupa 6844 (MO!, USM!); idem, 30 Mar 1939, C. Vargas 4505 (MO!); Urubamba, 23 Apr 1982, B. Peyton 51 (MO!); idem, 04 Jul 1982, B. Peyton 766 (MO!); idem, 17 Aug 1982, B. Peyton 1053 (GH!, MO!); idem, 05 Nov 1988, A. Tupayachi 758 (GH!, MO!, NY!); idem, 30 Dec 1963, L. Valenzuela 14983 (GH!). Huánuco: Yanano, 13-16 May 1923 J.F. Macbride 3826 (G!). La Libertad: Pataz, 24 Feb 1986, K. Young 2988 (NY); s.l., 23 Jun 1974, A. López M. 8129 (G!, MO!, NY!). Lambayeque: Ferreñafe, 9 Jun 2012, L. García Llatas s.n. (USM). San Martín: Mariscal Cáceres, Huicungo, 15 Jun 2001, B. León 5249 (NY!, USM).
Adetogramma chrysolepis (Hook.) T.E.Almeida. A, B Scanning electron micrography (SEM); magnification 75× and 1750×, respectively A Sori showing sporangia and paraphysis (the latter indicated by an arrow head) B Spores. Images from Almeida & Giacomin 3121 (BHCB). Scale bars: A = 500 μm, B = 20 μm.
Adetogramma chrysolepis (Hook.) T.E.Almeida. A Fertile and sterile fronds B Detail of abaxial surface of fertile frond, showing laminar scales and paraphyses C Juvenile sterile leaves. (A) and (B) from C. Martín 730 (SI), (C) from C. Martín 479 (SI).
This work was supported by FAPEMIG (Fundação de Amparo à Pesquisa do Estado de Minas Gerais through grant CRA-APQ 01599-10, CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) through grants 478723/2010-5 and 563568/2010, and for productivity grants, all attributed to AS, and the ATM “Biodiversité actuelle et fossile” of the Muséum national d’Histoire naturelle (2012) attributed to SH. CNPq also provided research scholarships to TEA (555226/2010-7 and 202160/2011-4). We thank the Muséum national d’Histoire naturelle de Paris, France (MNHN), the Service de Systématique Moléculaire (SSM - MNHN) and Céline Bonillo for the support and help. L.L. Giacomin for helping with fieldwork and for carefully reviewing the manuscript. A. Field, A. Smith, B. León, D. Sanín, E. Smidt, J. Prado, J.A.N. Batista, and V.A.O. Dittrich for critical reviews and comments. T. Martínez for guiding us in Bolivia; to the curators of all visited herbaria; Claudia Martín for the pictures, and J.T. Mickel for allowing the use of the drawings.
Collection information for voucher specimens and GenBank accession numbers for sequences used in this study. Locality and voucher specimen (herbarium) are given to sequences newly generated in this study. *sequences submitted to GenBank
|Adenophorus oahuensis (Copel.) L.E.Bishop||AY057382||AY096236|
|Adetogramma chrysolepis (Hook.) T.E.Almeida; Bolivia, Almeida 3083 (BHCB)||KY847865*||KY847858*|
|Adetogramma chrysolepis (Hook.) T.E.Almeida; Bolivia, Almeida 3121 (BHCB)||KY847859*||KY847863*|
|Aglaomorpha pilosa (J.Sm.) Copel.||AY529156||AY529180|
|Alansmia cultrata (Willd.) Moguel & M.Kessler||GU376496||JN654943|
|Arthromeris himalayensis (Hook.) Ching||JQ685378||JQ685442|
|Campyloneurum gracile A.Rojas; Panama, Salino 15357 (BHCB)||KY847860*||KY847864*|
|Campyloneurum phyllitidis (L.) C.Presl||KT780752||KT794132|
|Davallia solida (G.Forst.) Sw.||AY096193||AY096210|
|Dictymia brownii (Wikstr.) Copel.||DQ227292||DQ227295|
|Goniophlebium percussum (Cav.) W.H.Wagner & Grether||AY362561||AY362628|
|Gymnogrammitis dareiformis (Hook.) Tardieu & C.Chr.||AY096201||JQ685456|
|Lecanopteris carnosa (Reinw.) Blume||AF470322||AY096227|
|Lellingeria apiculata (Klotzsch) A.R.Sm. & R.C.Moran||GU387021||GU387046|
|Lellingeria brevistipes (Kuhn) A.R.Sm. & R.C.Moran||GU387030||GU387049|
|Lemmaphyllum carnosum (Hook.) C.Presl||GU126698||GU126717|
|Lepidomicrosorium subhemionitideum (Christ) P.S.Wang||GU126693||GU126711|
|Lepisorus heterolepis (Rosenst.) Ching||GQ256270||GQ256344|
|Leptochilus digitatus (Baker) Noot.||JX103695||EU363250|
|Leucotrichum madagascariense Rakotondr. & Rouhan||JN654924||JN654949|
|Leucotrichum organense (Gardner) Labiak||GU376490||JN654946|
|Loxogramme salicifolia (Makino) Makino||DQ227294||DQ227297|
|Melpomene flabelliformis (Poir.) A.R.Sm. & R.C.Moran||GU387028||GU387114|
|Melpomene melanosticta (Kunze) A.R.Sm. & R.C.Moran||GU387024||GU387115|
|Microgramma baldwinii Brade; Brazil, Almeida 2631 (BHCB)||KY847861*||KY847866*|
|Microgramma bifrons (Hook.) Lellinger||AY362582||AY362654|
|Microgramma mauritiana (Willd.) Tardieu||DQ642148||DQ642185|
|Microgramma persicariifolia (Schrad.) C.Presl||KT780753||KT794133|
|Microsorum membranaceum (D.Don) Ching||EU482963||AY725047|
|Mycopteris attenuatissima (Copel.) Sundue||GU476927||GU387121|
|Mycopteris subtilis (Klotzsch) Sundue||GU476875||GU387128|
|Neocheiropteris palmatopedata (Baker) Christ||JX103706||GQ256396|
|Niphidium crassifolium (L.) Lellingerl; Brazil, Almeida 3247 (BHCB)||KY847862*||KY847867*|
|Niphidium nidulare (Rosenst.) Lellinger||EF551064||EF551080|
|Oleandra pistillaris (Sw.) C.Chr||AB232405||AY096209|
|Pecluma ptilotos (Kunze) M.G.Price||AY362588||AY362661|
|Phlebodium pseudoaureum (Cav.) Lellinger||AY362589||AY362663|
|Platycerium andinum Baker||DQ164446||DQ164477|
|Pleopeltis angusta Willd.||EU650122||EU650161|
|Pleopeltis desvauxii (Klotzsch) Salino||AY362584||AY362657|
|Polypodium vulgare L.||JF832081||EF551081|
|Pyrrosia angustata (Sw.) Ching||DQ642165||DQ642204|
|Selliguea plantaginea Brack.||EF463262||EU128510|
|Serpocaulon articulatum (C.Presl) Schwartsb. & A.R.Sm.||DQ151910||DQ151935|
|Serpocaulon dissimile (L.) A.R.Sm.||DQ151908||DQ151933|
|Serpocaulon levigatum (Cav.) A.R.Sm.||DQ151917||EF551103|
|Serpocaulon loriceum (L.) A.R.Sm.||EF551074||EF551104|
|Stenogrammitis hellwigii (Mickel & Beitel) Labiak||GU386990||GU387058|
|Stenogrammitis hildebrandtii (Hieron.) Labiak||GU386975||GU387059|
|Synammia feuillei (Bertero) Copel.||AY362597||AY362670|