Research Article |
Corresponding author: Yao-Moan Huang ( huangym@tfri.gov.tw ) Academic editor: Thais Almeida
© 2017 Cheng-Wei Chen, Michael Sundue, Li-Yaung Kuo, Wei-Chih Teng, Yao-Moan Huang.
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:
Chen C-W, Sundue M, Kuo L-Y, Teng W-C, Huang Y-M (2017) Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae. PhytoKeys 78: 83-107. https://doi.org/10.3897/phytokeys.78.12040
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The monotypic fern genus Dryopolystichum Copel. combines a unique assortment of characters that obscures its relationship to other ferns. Its thin-walled sporangium with a vertical and interrupted annulus, round sorus with peltate indusium, and petiole with several vascular bundles place it in suborder Polypodiineae, but more precise placement has eluded previous authors. Here we investigate its phylogenetic position using three plastid DNA markers, rbcL, rps4-trnS, and trnL-F, and a broad sampling of Polypodiineae. We also provide new data on Dryopolystichum including spore number counts, reproductive mode, spore SEM images, and chromosome counts. Our maximum-likelihood and Bayesian-inference phylogenetic analyses unambiguously place Dryopolystichum within Lomariopsidaceae, a position not previously suggested. Dryopolystichum was resolved as sister to a clade comprising Dracoglossum and Lomariopsis, with Cyclopeltis as sister to these, but clade support is not robust. All examined sporangia of Dryopolystichum produced 32 spores, and the chromosome number of sporophyte somatic cells is ca. 164. Flow cytometric results indicated that the genome size in the spore nuclei is approximately half the size of those from sporophyte leaf tissues, suggesting that Dryopolystichum reproduces sexually. Our findings render Lomariopsidaceae as one of the most morphologically heterogeneous fern families. A recircumscription is provided for both Lomariopsidaceae and Dryopolystichum, and selected characters are briefly discussed considering the newly generated data.
Fern, morphology, Papua New Guinea, phylogeny, recircumscription, taxonomy, the Solomon Islands
Dryopolystichum Copel., with its single species D. phaeostigma (Ces.) Copel., is distributed along streams in lowland forests in New Guinea, the Bismarck Archipelago, and the Solomon Islands (
Although Copeland did not provide an etymological explanation, the name Dryopolystichum presumably reflects the combination of peltate indusium (which is similar to those of polystichoid ferns) and pinnate-pinnatifid lamina division (which is similar to that of most Dryopteris). Such a combination of characters resulted in taxonomic confusion giving that peltate indusia are never found in Dryopteris, and the laminae of Dryopolystichum do not include prominulous segment apices, the hallmark of polystichoid ferns (
Despite recent advances in fern phylogenetics and classification, the position of Dryopolystichum remains unclear. The thin-walled sporangium with a vertical and interrupted annulus, round sorus, and petiole with several vascular bundles suggest that this genus belongs to suborder Polypodiineae (= eupolypods I) (
One other conspicuous character of Dryopolystichum not emphasized by previous authors is that the distal pinnae are decurrent onto the rachis, and the basal pinnules of its distal pinnae are served by veins that emerge from the rachis, rather than the pinna costa (Fig.
Subsequent to its establishment as a new genus in Genera Filicum (
To resolve the phylogenetic placement of Dryopolystichum, we employ a molecular phylogenetic approach using three chloroplast DNA regions, rbcL, rps4-trnS, and trnL-F. Based on our observations, we further provide new data on Dryopolystichum including spore counts, reproductive mode, spore SEM images, and a chromosome count. Finally, we discuss its diagnostic characters in the light of the inferred phylogeny.
We examined the morphology of Dryopolystichum phaeostigma using material collected from the Solomon Islands (Braithwaite R.S.S.4557, SING; SITW10443, BSIP, TAIF, TNM) and Papua New Guinea (James & Sundue 1688, BISH, LAE, VT).
Living plants of SITW10443 were transplanted to the Dr. Cecilia Koo Botanic Conservation Center in Taiwan (KBCC). The collection of SITW10443 was made under the “Census and Classification of Plant Resources in the Solomon Islands” project (http://siflora.nmns.edu.tw/). Mitotic chromosomes were counted from these cultivated plants following the protocol of
Fertile pinnae of SITW10443 were air-dried in an envelope for one day to release the spores. The spores were observed and measured by a tabletop scanning electron microscope (TM-3000 Hitachi, Ibaraki, Japan). The sizes (the length of equatorial axes including the perine ornamentation) of 35 randomly selected spores were measured. Five intact sporangia were observed under a stereo microscope (Leica MZ6, Wetzlar, Germany) to count the number of spores per sporangium.
The genome sizes of spore and leaf nuclei of SITW10443 were examined by flow cytometry in order to infer the reproductive mode (
Total DNA was extracted using a modified CTAB-Qiagen column protocol (
The PCR amplifications were performed in 16 μl reactions containing ca. 10 ng template DNA, 1×Taq DNA Polymerase Master Mix RED solution (Ampliqon, Denmark), and 1 μl each of 10 μM primers. The PCR reactions were carried out in a GeneAmp PCR System 9700 (Applied Biosystems, Carlsbad, California, USA). Thermocycling conditions were the same for PCRs of these three regions and comprised an initial denaturation of 2 minutes at 94°C followed by a core sequence of 35 repetitions of 94°C for 1 minute, 55°C for 1 minute, and 72°C for 1 minute followed by a final extension of 10 minutes at 72°C. Resulting PCR products were sequenced using the same PCR primers with BigDyeTM terminator (Applied Biosystems, Carlsbad, California, USA). The newly generated sequences were deposited in GenBank. GenBank accession numbers and voucher information are provided in Appendix.
Initial BLAST against the NCBI nucleotide database (
Sequences were aligned using Geneious v6.1.8 (
We addressed the possibility of phylogenetic bias due to long branches following the recommendation of Siddal and Whiting (1999). Since Dracoglossum and Lomariopsis were resolved on long branches in preliminary analyses (not shown), we conducted two additional analyses in which each one of the two long-branched genera, Dracoglossum and Lomariopsis, was excluded to examine whether phylogenetic placement and branch support for Dryopolystichum’s placement changed. Since maximum parsimony (MP) phylogeny is considered to be more susceptible to long-branch attraction (
All single-region phylogenies resolved Dryopolystichum phaeostigma in Lomariopsidaceae, but with two slightly different topologies. The rbcL and rps4-trnS phylogenies placed D. phaeostigma sister to a clade of Dracoglossum + Lomariopsis with 93% and 72% maximum likelihood bootstrap percentages (BS), respectively (Suppl. materials
Simplified maximum likelihood phylogram of Polypodiineae obtained from the rbcL + rps4-trnS + trnL-F combined dataset. Maximum likelihood bootstrap percentages (BS) are provided at each node. Thickened lines indicate Bayesian inference posterior probability (PP) ≥ 0.9. Original phylogram with support values for all the nodes is available in Suppl. materials
Removing Dracoglossum from the analysis had little effect on the topology within Lomariopsidaceae, and BS supports for the generic placement of Dryopolystichum remained low (≤ 70%, data not shown). In contrast, the removal of Lomariopsis resulted in higher BS values for all clades within Lomariopsidaceae (≥ 99%, data not shown). MP analyses also resulted in a clade comprising all the Lomariopsidaceae genera and Dryopolystichum, but Dryopolystichum was resolved as sister to Cyclopeltis (data not shown).
All examined sporangia (SITW10443) produced 32 normal spores, and the mean spore length was 64.1 ± 4.5 μm (Fig.
The reconstructed maximum likelihood and Bayesian inference phylogenies unambiguously resolved Dryopolystichum within Lomariopsidaceae (Fig.
Phylogenetic analyses using DNA sequences have served as the basis for redrawing fern classifications in the 21th century (
Subsequent molecular phylogenetic analyses demonstrated that most genera previously treated in Lomariopsidaceae should be transferred to Dryopteridaceae (
More recently, the neotropical genus Dracoglossum was established (
Comparison of morphological characters of the five Lomariopsidaceae genera [based on
Genera | Cyclopeltis | Dracoglossum | Dryopolystichum | Lomariopsis | Thysanosoria |
---|---|---|---|---|---|
Habit | terrestrial | terrestrial | terrestrial | hemiepiphyte | hemiepiphyte |
Rhizome | erect | short creeping | erect | climbing | climbing |
Frond division* | pinnate | simple | pinnate-pinnatifid | pinnate | pinnate |
Pinnae articulation | articulate | – | not articulate | articulate | articulate |
Venation | free | reticulate, with included veinlet | free | free | free |
Rachis-costa architecture | prominent | prominent | grooved | grooved or flat | grooved |
Sporangia | form rounded sori | form rounded sori | form rounded sori | acrostichoid | form rounded sori |
Indusia | peltate if present | peltate if present | peltate | absent | absent |
Perine ornamentation | broad folds | narrow crests | narrow crests | various | broad folds |
Lomariopsis Fée, Mém. Foug., 2. Hist. Acrostich.: 10. 1845.
Habit erect, creeping, or climbing; rhizomes dictyostelic, the ventral meristele elongate in transverse section or not; scaly at least when young; scales non-clathrate, basally attached or shallowly peltate, margins entire, toothed, or ciliate; fronds monomorphic or dimorphic; petioles with multiple vascular bundles arranged in a U-shape; laminae simple, pinnate, or pinnate-pinnatifid, provided distally with proliferous buds or not; pinnae articulate to the rachis or not; veins free, ± parallel or pinnate; sori acrostichoid or discrete and then round, with peltate indusia or exindusiate; spores brown, olive or green, chlorophyllous or not, bilateral, monolete, perine loosely attached, variously winged or ornamented.
Five genera and an estimated 70 species. Thysanosoria is included based on its morphological similarity to Lomariopsis (
Dryopolystichum phaeostigma (Ces.) Copel., Gen. Fil. 125, t. 4. 1947.
Habit terrestrial, on slopes along streams at lowland forests; rhizome short erect, stout and woody, apex densely scaly, blackish sclerenchyma strands visible in sections; scales dark brown, linear-lanceolate, entire, not clathrate; fronds approximate, stipe not articulate, scaly at base, scales similar to those on rhizome; lamina ovate, pinnate-pinnatifid, catadromous, subleathery, nearly glabrous, only very sparse narrow scales on rachis, costa, and costule; rachis and costa grooved adaxially, not connected to each other; veins free, pinnate, veins of basal pinnules on upper pinnae emerge from the rachis rather than costa, all veins terminating in a prominent hydathode, not reaching frond margin; sori round, dorsally on veinlets near hydathode, indusiate; indusia round, persistent, superior, entire, brownish, thick; sporangia long-stalked, annulus with ca. 14 indurated cells, 32 normal spores in each sporangium; spores monolete, 64.1 ± 4.5 μm in lateral view, surface with broadly winged wall; 2n = ca. 164.
Monotypic.
Aspidium phaeostigma Ces., Rend. Ac. Napoli 16: 26, 29. 1877.
Type. Papua New Guinea. Andai, Beccari 12533 (FI [FI013622]).
Dryopteris phaeostigma (Ces.) C.Chr., Index Filic. 284. 1905
Type. Based on Aspidium phaeostigma Ces.
Dryopteris tamatana C.Chr., Index Filic., Suppl. (1906-1912) 40. 1913.
Replaced: Dryopteris kingii Copel., Phillipp. J. Sci., C 6: 73. 1911., not Dryopteris kingii (Bedd.) C.Chr., Index Filic. 273. 1905.
Type. Papua New Guinea. Tamata, C. King 149 (MICH [MICH1287049]).
Polystichum lastreoides Rosenst., Repert. Spec. Nov. Regni Veg. 9: 425. 1911.
Type. Papua New Guinea. C. King 194 (MICH [MICH1190927]).
Dryopteris ledermannii Brause, Bot. Jahrb. Syst. 56: 90. 1920.
Type. Papua New Guinea. Sepik, Ledermann 9619 (B [B_20_005865], L [L0063060], S [S-P-8581]).
Dryopteris cyclosorus Alderw., Nova Guinea 14: 21. 1924.
Type. Indonesia. Irian Jaya, H. J. Lam 1086 (BO [BO1529719, BO1529720], K [K000666126], L [L0051583], U [U0007385]).
Based on Aspidium phaeostigma Ces.
Equal to the genus.
New Guinea, the Bismark archipelago, and the Solomon Islands.
Perine architecture of Dryopolystichum is very similar to that of Dracoglossum plantagineum (
Blackish sclerenchyma strands are visible in the rhizome sections of Dryopolystichum (Fig.
The rachis-costae architecture of Dryopolystichum is characterized by an adaxially sulcate rachis with grooves that do not connect to those of the pinna-costae. The rachis is also narrowly winged laterally. Both characters are seen in Thysanosoria and in some species of Lomariopsis (
The chromosome number in somatic cells of Dryopolystichum phaeostigma was ca. 164 (Fig.
Our flow cytometry and spore count results indicate that Dryopolystichum phaeostigma is sexually reproducing and has 32 spores per sporangium (Fig.
We have shown, based on molecular phylogenetic evidence, the placement of Dryopolystichum within Lomariopsidaceae. A revised description was provided for both Lomariopsidaceae and Dryopolystichum resulting from a review of literature and our own observations. Future studies using an expanded dataset are necessary to resolve intergeneric relationships in Lomariopsidaceae.
We are grateful to Kathleen Pryer’s lab for sharing the material of Dracoglossum. Peter Hovenkamp, Thais Almeida, David Barrington, and two anonymous reviewers provided valuable comments on an earlier draft of this manuscript. The curators and staffs of herbaria BSIP, SING, TAIF, and VT for providing access to their collections. We also thank Robbin Moran and Wita Wardani for checking specimens at NY and BO, respectively. Field work in Solomon Islands was supported by Taiwan International Cooperation and Development Fund (TH410-2012-085), Taiwan Forestry Research Institute (102AS-4.1.1-FI-G1), and Dr. Cecilia Koo Botanic Conservation Center (KBCC) for CWC.
Individuals sampled in this study. For each individual, the species name and GenBank accession numbers (rbcL, rps4-trnS, trnL-F) are provided. A n-dash (–) indicates unavailable information; new sequences are in bold.
Taxon | Genbank accession numbers | ||
---|---|---|---|
rbcL | rps4-trnS | trnL-F | |
Dryopteridaceae | |||
Arachniodes aristata (G.Forst.) Tindale | KJ464418 | – | KJ464592 |
Arachniodes denticulata (Sw.) Ching | KJ464419 | – | KJ464593 |
Arthrobotrya articulata J.Sm. | – | GU376714 | GU376565 |
Arthrobotrya wilkesiana Copel. | – | GU376719 | GU376569 |
Bolbitis acrostichoides (Afzel.) Ching | KJ464420 | GU376644 | GU376500 |
Bolbitis aliena (Sw.) Alston | – | GU376646 | GU376502 |
Bolbitis angustipinna (Hayata) H.Ito | – | GU376654 | GU376509 |
Bolbitis appendiculata (Willd.) K.Iwats. | – | GU376647 | GU376503 |
Bolbitis auriculata (Lam.) Alston | KJ464421 | GU376649 | GU376505 |
Bolbitis bipinnatifida (J.Sm.) K.Iwats. | – | GU376676 | GU376530 |
Bolbitis fluviatilis (Hook.) Ching | – | GU376656 | GU376510 |
Bolbitis gemmifera (Hieron.) C.Chr. | – | GU376657 | GU376511 |
Bolbitis heteroclita (Pr.) Ching | – | GU376659 | GU376513 |
Bolbitis heudelotii (Bory) Alston | – | GU376662 | GU376515 |
Bolbitis humblotii (Baker) Ching | KJ464422 | GU376663 | GU376516 |
Bolbitis lonchophora (Kunze) C.Chr. | – | GU376664 | GU376517 |
Bolbitis major (Bedd.) Hennipman | – | GU376665 | GU376518 |
Bolbitis portoricensis (Sprengel) Hennipman | – | GU376670 | GU376523 |
Bolbitis salicina (Hook.) Ching | – | GU376671 | GU376525 |
Bolbitis semipinnatifida (Fée) Alston | – | GU376672 | GU376526 |
Bolbitis serratifolia (Mertens) Schott | – | GU376673 | GU376527 |
Bolbitis sinuata (C.Presl) Hennipman | – | GU376675 | GU376529 |
Bolbitis tibetica Ching & S.K.Wu | – | GU376677 | GU376531 |
Ctenitis eatonii (Baker) Ching | KF709483 | – | KJ196645 |
Ctenitis sinii (Ching) Ohwi | – | – | KJ196643 |
Ctenitis subglandulosa (Hance) Ching | – | – | KJ196655 |
Ctenitis yunnanensis Ching & Chu H.Wang | – | – | KJ196715 |
Cyclodium heterodon var. heterodon | KJ464425 | – | KJ464596 |
Cyclodium rheophilum A.R.Sm. | KJ464426 | – | KJ464597 |
Dryopteris apiciflora (Wall. ex Mett.) Kuntze | – | – | KJ196641 |
Dryopteris christensenae (Ching) Li Bing Zhang | – | – | KJ196679 |
Dryopteris heterolaena C.Chr. | – | – | KJ196623 |
Dryopteris integriloba C.Chr. | – | – | KJ196701 |
Dryopteris mariformis Rosenst. | – | – | KJ196686 |
Dryopteris nidus (Baker) Li Bing Zhang | – | – | KJ196687 |
Dryopteris patula (Sw.) Underw. | KJ464427 | – | KJ464598 |
Dryopteris polita Rosenst. | – | – | KJ196700 |
Dryopteris squamiseta (Hook.) Kuntze | – | GU376678 | KJ196632 |
Dryopteris wallichiana (Spreng.) Hyl. | KJ464428 | GU376680 | KJ464599 |
Elaphoglossum amygdalifolium (Mett.) Christ | – | GU376681 | – |
Elaphoglossum burchellii (Baker) C.Chr. | – | GU376682 | GU376533 |
Elaphoglossum decoratum (Kunze) T.Moore | KJ464429 | GU376683 | KJ464600 |
Elaphoglossum guentheri Rosenst. | – | GU376684 | GU376535 |
Elaphoglossum langsdorffii T.Moore | – | GU376685 | GU376536 |
Elaphoglossum lloense (Hook.) T.Moore | – | GU376686 | GU376537 |
Elaphoglossum luridum Christ | – | – | GU376538 |
Elaphoglossum squamipes (Hook.) T.Moore | – | – | GU376539 |
Lastreopsis amplissima (C.Presl) Tindale | KJ464432 | – | KJ464604 |
Lastreopsis decomposita (R.Br.) Tindale | KJ464439 | – | – |
Lastreopsis hispida (Sw.) Tindale | KJ464446 | – | KJ464614 |
Lastreopsis killipii (C.Chr. & Maxon) Tindale | KJ464448 | KF709505 | – |
Lastreopsis marginans (F.Muell.) Tindale | KJ464449 | GU376691 | KJ464616 |
Lastreopsis poecilophlebia (Hook.) Labiak, Sundue & R.C.Moran | KJ464423 | GU376692 | KJ464594 |
Lastreopsis tenera (R.Br.) Tindale | KJ464467 | GU376699 | KJ464636 |
Lastreopsis tripinnata (F.Muell. ex Benth.) Labiak, Sundue & R.C.Moran | KJ464491 | GU376700 | – |
Lastreopsis walleri Tindale | KJ464472 | GU376701 | – |
Lastreopsis wurunuran (Domin) Tindale | KJ464474 | GU376704 | – |
Lomagramma brooksii Copel. | – | GU376705 | GU376542 |
Lomagramma cordipinna Holttum | – | GU376707 | GU376543 |
Lomagramma lomarioides (Blume) J.Sm. | – | – | GU376550 |
Lomagramma matthewii (Ching) Holttum | KJ464476 | – | KJ464640 |
Lomagramma perakensis Bedd. | – | – | GU376552 |
Lomagramma pteroides J.Sm. | – | – | GU376555 |
Lomagramma sinuata C.Chr. | – | – | GU376556 |
Lomagramma sumatrana Alderw. | – | – | GU376558 |
Maxonia apiifolia (Sw.) C.Chr. | KJ464477 | GU376709 | KJ464641 |
Megalastrum abundans (Rosenst.) A.R.Sm. & R.C.Moran | KJ464478 | – | KJ464642 |
Megalastrum atrogriseum (C.Chr.) A.R.Sm. & R.C.Moran | KJ464479 | GU376710 | KJ464643 |
Megalastrum connexum (Kaulf.) A.R.Sm. & R.C.Moran | KJ464481 | – | KJ464645 |
Megalastrum lanatum (Fée) Holttum | KJ464483 | – | KJ464647 |
Megalastrum littorale R.C.Moran, J.Prado & Labiak | – | GU376651 | GU376561 |
Megalastrum macrotheca (Fée) A.R.Sm. & R.C.Moran | KJ464484 | GU376697 | KJ464648 |
Megalastrum vastum (Kunze) A.R.Sm. & R.C.Moran | KJ464487 | GU376658 | KJ464651 |
Mickelia bernoullii (Kuhn ex Christ) R.C.Moran, Labiak & Sundue | – | GU376666 | GU376506 |
Mickelia guianensis (Aubl.) R.C.Moran, Labiak & Sundue | – | GU376667 | GU376548 |
Mickelia hemiotis (Maxon) R.C.Moran, Labiak & Sundue | – | – | GU376512 |
Mickelia nicotianifolia (Sw.) R.C.Moran, Labiak & Sundue | – | KF667557 | GU376519 |
Mickelia oligarchica (Baker) R.C.Moran, Labiak & Sundue | KJ464489 | – | GU376520 |
Mickelia scandens (Raddi) R.C. Moran, Labiak & Sundue | – | GU376696 | GU376547 |
Olfersia cervina Kunze | KJ464493 | DQ153079 | KJ464652 |
Parapolystichum acuminatum (Houlston) Labiak, Sundue & R.C.Moran | KJ464430 | KC977454 | KJ464601 |
Parapolystichum boivinii (Baker) Rouhan | KJ464435 | – | KJ464607 |
Parapolystichum confine (Maxon ex C.Chr.) Labiak, Sundue & R.C.Moran | KJ464438 | – | – |
Parapolystichum effusum (Sw.) Ching | KJ464441 | – | – |
Parapolystichum effusum subsp. divergens (Willd. ex Schkuhr) Tindale | KJ464440 | – | – |
Parapolystichum excultum (Mett.) Labiak, Sundue & R.C.Moran | – | KF709501 | GU376541 |
Parapolystichum glabellum (A.Cunn.) Labiak, Sundue & R.C.Moran | KJ464445 | KF709503 | KJ464613 |
Parapolystichum microsorum (Endl.) Labiak, Sundue & R.C.Moran | KJ464451 | GU376712 | KJ464617 |
Parapolystichum perrierianum (C.Chr.) Rouhan | KJ464455 | – | KJ464623 |
Parapolystichum rufescens (Blume) Labiak, Sundue & R.C.Moran | KJ464461 | – | KJ464629 |
Parapolystichum vogelii (Hook.) Rouhan | KJ464470 | ||
Parapolystichum windsorensis (D.L.Jones & B.Gray) Labiak, Sundue & R.C.Moran | KJ464473 | – | KJ464639 |
Pleocnemia conjugata C.Presl | – | GU376713 | KF709510 |
Pleocnemia cumingiana C.Presl | KJ196828 | – | KJ196705 |
Pleocnemia dahlii (Hieron.) Holttum | KJ196829 | – | KJ196706 |
Pleocnemia hemiteliiformis (Racib.) Holttum | KF709482 | KF667560 | KF709511 |
Pleocnemia irregularis (C.Presl) Holttum | KF709491 | – | KF709513 |
Pleocnemia leuzeana (Gaudich.) C.Presl | KJ196830 | – | – |
Pleocnemia olivacea (Copel.) Holttum | KJ464495 | – | – |
Pleocnemia presliana Holttum | KJ464496 | KF667561 | – |
Pleocnemia rufinervis Nakai | JF303976 | KF667562 | – |
Pleocnemia winitii Holttum | EF460686 | – | KF709515 |
Polybotrya alfredii Brade | KJ464497 | KF667563 | KJ464653 |
Polybotrya andina C.Chr. | KJ464498 | KP271084 | KJ464654 |
Polybotrya pubens Mart. | KJ464499 | KP271085 | – |
Polystichum tsus-simense var. mayebarae (Tagawa) Sa.Kurata | AB575224 | – | DQ150408 |
Pseudotectaria biformis (Mett.) Holttum | – | – | KF897951 |
Pseudotectaria decaryana (C.Chr.) Tardieu | – | – | KF897952 |
Rumohra adiantiformis (G.Forst.) Ching | KJ464500 | – | KJ464655 |
Rumohra berteroana (Colla) J.J. Rodr. | KJ464503 | – | KJ464657 |
Stigmatopteris ichthiosma (Sodiro) C.Chr. | KJ464504 | – | KJ464658 |
Stigmatopteris killipiana Lellinger | KJ464505 | – | KJ464659 |
Stigmatopteris lechleri (Mett) C.Chr. | KJ464506 | KP271087 | KJ464660 |
Stigmatopteris sordida (Maxon) C.Chr. | KJ464507 | – | KJ464661 |
Teratophyllum koordersii Holttum | – | – | GU376566 |
Teratophyllum ludens (Fée) Holttum | – | – | GU376567 |
Teratophyllum wilkesianum Holttum | KJ464508 | – | – |
Nephrolepidaceae | |||
Nephrolepis abrupta (Bory) Mett. | HM748137 | KF667559 | – |
Nephrolepis acutifolia (Desv.) Christ. | HM748139 | – | – |
Nephrolepis biserrata (Sw.) Schott | AB575227 | GU376688 | – |
Nephrolepis brownii (Desv.) Hovenkamp & Miyam. | KR816691 | – | – |
Nephrolepis cordifolia (L.) C.Presl | AB575228 | – | – |
Nephrolepis davalliae Alderw. | HM748147 | – | – |
Nephrolepis davallioides Kunze | HM748148 | GU376690 | – |
Nephrolepis exaltata (L.) Schott | HM748149 | – | – |
Nephrolepis falcata (Cav.) C.Chr. | HM748150 | – | – |
Nephrolepis falciformis J.Sm. | AB232404 | – | – |
Nephrolepis lauterbachii (Christ) Christ | HM748153 | – | – |
Nephrolepis pectinata (Willd.) Schott | HM748155 | – | – |
Nephrolepis pendula (Raddi) J.Sm. | HM748156 | – | – |
Nephrolepis radicans (Burm.) Kuhn | HM748157 | – | – |
Nephrolepis rivularis (Vahl) Mett. | HM748158 | – | – |
Nephrolepis undulata J.Sm. | HM748159 | – | – |
Lomariopsidaceae | |||
Cyclopeltis crenata (Fée) C.Chr. | DQ054517 | EF540718 | DQ51448 |
Cyclopeltis novoguineensis Rosenst. | KY397974 | KY397978 | KY397970 |
Cyclopeltis semicordata (Sw.) J.Sm. | EF463234 | KY397977 | KY397969 |
Dracoglossum plantagineum (Jacq.) Christenh. | KC914564 | KY397979 | KY397971 |
Dracoglossum sinuatum (Fée) Christenh. | – | – | KU605106 |
Dryopolystichum phaeostigma (Ces.) Copel. | KY397972 | KY397976 | KY397968 |
Lomariopsis crassifolia Holttum | – | – | DQ396559 |
Lomariopsis guineensis (Underw.) Alston | – | KJ628952 | DQ396560 |
Lomariopsis hederacea Alston | – | – | DQ396561 |
Lomariopsis jamaicensis (Underw.) Holttum | – | – | DQ396562 |
Lomariopsis japurensis (C.Martius) J.Sm. | – | – | DQ396563 |
Lomariopsis kunzeana (Underw.) Holttum | – | – | DQ396569 |
Lomariopsis latipinna Stolze | – | – | DQ396571 |
Lomariopsis lineata (C.Presl) Holttum | – | – | DQ396572 |
Lomariopsis longicaudata (Bonap.) Holttum | – | – | Q396573 |
Lomariopsis madagascarica (Bonap.) Alston | – | – | DQ396575 |
Lomariopsis mannii (Underw.) Alston | – | – | DQ396577 |
Lomariopsis marginata (Schrad.) Kuhn | AY818677 | – | DQ396578 |
Lomariopsis maxonii (Underw.) Holttum | – | – | DQ396580 |
Lomariopsis muriculata Holttum | – | – | DQ396582 |
Lomariopsis palustris (Hook.) Mett. ex Kuhn | – | HM748162 | DQ396585 |
Lomariopsis pervillei Kuhn | – | – | DQ396586 |
Lomariopsis pollicina (Willemet) Mett. ex Kuhn | EF463235 | – | DQ396588 |
Lomariopsis prieuriana Fée | – | – | DQ396590 |
Lomariopsis recurvata Fée | – | – | DQ396592 |
Lomariopsis rossii Holttum | – | – | DQ396594 |
Lomariopsis salicifolia (Kunze) Lellinger | – | – | DQ396595 |
Lomariopsis sorbifolia (L.) Fée | EF463236 | – | – |
Lomariopsis spectabilis Mett. | AB232401 | – | KJ196685 |
Lomariopsis vestita E.Fourn. | – | – | DQ396598 |
Lomariopsis wrightii Mett. | – | – | DQ396600 |
Tectariaceae | |||
Arthropteris altescandens J.Sm. | KF667636 | KF667550 | KF667606 |
Arthropteris articulata (Brack.) C.Chr. | KC977367 | KC977437 | KC977411 |
Arthropteris beckleri (Hook.) Mett. | U05605 | – | KF667607 |
Arthropteris cameroonensis Alston | KF667638 | – | – |
Arthropteris guinanensis H.G.Zhou & Y.Y.Huang | KC977364 | KC977442 | KC977404 |
Arthropteris monocarpa (Cordem.) C.Chr. | HM748132 | – | KF897941 |
Arthropteris orientalis (Gmel.) Posth. | HM748133 | KC977435 | KC977420 |
Arthropteris palisotii (Desv.) Alston | AB575230 | KC977427 | KC977406 |
Arthropteris parallela (Baker) C.Chr. | EF463266 | KC977453 | KC977425 |
Arthropteris paucivenia (C.Chr.) H.M.Liu, Hovenkamp & H.Schneid. | EF463268 | – | KC977426 |
Arthropteris repens (Brack.) C.Chr. | KC977368 | KC977438 | KC977412 |
Arthropteris tenella (G.Forst.) J.Sm. ex Hook.f. | KC977363 | KF011547 | KC977424 |
Hypoderris brauniana (H.Karst.) F.G.Wang & Christenh. | KF667647 | – | KF667618 |
Hypoderris brownii J.Sm. | KF667642 | – | KF667611 |
Hypoderris nicotianifolia (Baker) R.C.Moran, Labiak & J.Prado | KF667653 | – | KF667626 |
Pteridrys australis Ching | KJ196892 | – | KJ196678 |
Pteridrys cnemidaria (Christ) C.Chr. & Ching | KF709488 | – | KF709517 |
Pteridrys lofouensis (Christ) C.Chr. & Ching | EF460687 | KF667566 | – |
Pteridrys microthecia (Fée) C.Chr. & Ching | KJ196848 | – | KF709518 |
Pteridrys syrmatica (Willd.) C.Chr. & Ching | KJ196875 | – | KF709519 |
Tectaria acerifolia R.C.Moran | KF887170 | – | KF897954 |
Tectaria angulata (Willd.) Copel. | KJ196876 | – | KJ196656 |
Tectaria aurita (Sw.) S.Chandra | KJ196849 | – | KJ196631 |
Tectaria barberi (Hook.) Copel. | KJ196846 | – | KJ196628 |
Tectaria borneensis S.Y.Dong | KJ196854 | KF667555 | KJ196642 |
Tectaria cicutaria (L.) Copel. | KF667649 | – | KF667620 |
Tectaria coadunata (J.Sm.) C.Chr. | KJ196851 | – | KJ196661 |
Tectaria crenata Cav. | KF667650 | KF667568 | KF667621 |
Tectaria decurrens (C.Presl) Copel. | AB575232 | – | DQ514524 |
Tectaria devexa (Kunze ex Mett.) Copel. | AB575233 | KP271088 | KF897956 |
Tectaria dilacerata (Kunze) Maxon | KF887173 | – | KF897957 |
Tectaria fauriei Tagawa | AB575234 | – | KJ196658 |
Tectaria fernandensis C.Chr. | KF887174 | – | KF897958 |
Tectaria gigantea (Blume) Copel. | KJ196853 | – | KJ196660 |
Tectaria griffithii (Baker) Ching | KF667652 | – | KF667624 |
Tectaria grossedentata Ching & Chu H.Wang | KJ196882 | KP271089 | KJ196667 |
Tectaria harlandii (Hook.) C.M.Kuo | AB575231 | – | KJ196648 |
Tectaria harlandii (Hook.) C.M.Kuo | KF887178 | – | KF897961 |
Tectaria heracleifolia (Willd.) Underw. | KF887180 | – | KF897963 |
Tectaria herpetocaulos Ching & Chu H. Wang | KJ196884 | – | KJ196669 |
Tectaria heterocarpa C.V.Morton | KF887181 | – | KF897964 |
Tectaria impressa (Fée) Holttum | KJ196841 | – | KF897965 |
Tectaria kusukusensis (Hayata) Lellinger | EF460681 | – | KF897968 |
Tectaria labrusca (Hook.) Copel. | KJ196818 | – | KJ196692 |
Tectaria luchunensis S.K.Wu | KJ196845 | KP271090 | KJ196627 |
Tectaria macleanii (Copel.) S.Y.Dong | KJ196810 | – | KJ196680 |
Tectaria melanocaula (Blume) Copel. | KJ196832 | – | KJ196709 |
Tectaria morsei (Baker) P.J.Edwards ex S.Y.Dong | KJ196893 | KF667570 | KF561675 |
Tectaria nayarii Mazumdar | EF463267 | – | KJ196699 |
Tectaria paradoxa (Fée) Sledge | KF887189 | – | KF897971 |
Tectaria phaeocaulis (Rosenst.) C.Chr. | AB232397 | KF709499 | KF897972 |
Tectaria pica (L.) C.Chr. | KF887191 | GU376715 | KF897973 |
Tectaria polymorpha (Wall. ex Hook.) Copel. | KJ196888 | GU376716 | KJ196657 |
Tectaria prolifera (Hook.) R.M.Tryon & A.F.Tryon | EF463273 | – | KF897974 |
Tectaria psomiocarpa S.Y.Dong | KJ196822 | KF667572 | KJ196698 |
Tectaria pubens R.C.Moran | KF887193 | KF667573 | KF897975 |
Tectaria quinquefida (Baker) Ching | KJ196885 | – | KJ396622 |
Tectaria repanda (Willd.) Holttum | KJ196831 | – | KJ196707 |
Tectaria sagenioides (Mett.) Christenh. | KF887194 | KF667575 | KF561672 |
Tectaria semipinnata (Roxb.) Morton | KJ196817 | KF667577 | KJ196691 |
Tectaria simonsii (Baker) Ching | AB575236 | – | KF897977 |
Tectaria singaporiana (Wall. ex Hook. & Grev.) Ching | KF887196 | – | KF897978 |
Tectaria subglabra (Holttum) S.Y.Dong | – | – | KJ196676 |
Tectaria subsageniacea (Christ) Christenh. | KF887197 | KF667576 | KF561670 |
Tectaria subtriphylla (Hook. & Arn.) Copel. | AB575237 | – | KF897980 |
Tectaria tricuspis (Bedd.) Copel. | KJ196820 | – | KJ196694 |
Tectaria variolosa (Wall. ex Hook.) C.Chr. | EF460690 | – | KF897982 |
Tectaria vasta (Blume) Copel. | KF667655 | – | KF667628 |
Tectaria vivipara Jermy & T.G.Walker | KF887201 | – | KF897983 |
Tectaria zeilanica (Houtt.) Sledge | AB232395 | – | KF709521 |
Triplophyllum crassifolium Holttum | KF887203 | – | KF897985 |
Triplophyllum fraternum (Mett.) Holttum | KF667657 | – | KF667630 |
Triplophyllum funestum (Kunze) Holttum | EF463276 | – | KF667631 |
Triplophyllum glabrum J.Prado & R.C.Moran | KF887207 | – | KF897989 |
Triplophyllum heudelotii Pic.Serm. | – | – | KF897990 |
Triplophyllum jenseniae (C.Chr.) Holttum | KF667660 | – | KF667633 |
Triplophyllum pentagonum (Bonap.) Holttum | KF667662 | – | KF667635 |
Triplophyllum pilosissimum (J.Sm. ex T.Moore) Holttum | – | – | KU605127 |
Triplophyllum securidiforme (Hook.) Holttum | – | – | KU605128 |
Triplophyllum vogelii (Hook.) Holttum | KF667661 | – | KF667634 |
Oleandraceae | |||
Oleandra articulata (Sw.) C.Presl | KF667644 | KF709500 | KF667613 |
Oleandra cumingii J.Sm. | KJ196816 | – | KJ196690 |
Oleandra neriiformis Cav. | KJ196815 | – | KJ196689 |
Oleandra pilosa Hook. | KF667646 | – | KF667615 |
Davalliaceae | |||
Davallodes hirsuta (J.Sm.) Copel. | AY096196 | – | – |
Davallodes yunnanensis (Christ) M.Kato & Tsutsumi | JX103718 | KC914565 | – |
Polypodiaceae | |||
Campyloneurum minus Fée | KF667665 | – | – |
Microgramma lycopodioides (L.) Copel. | KF667664 | – | – |
Niphidium longifolium (Cav.) C.V.Morton & Lellinger | KF667663 | KF709495 | – |
Figure
Data type: statistical data
Explanation note: Maximum likelihood bootstrap percentages (BS) are provided at each node. Thickened lines indicate Bayesian inference posterior probability (PP) ≥ 0.9.
Figure
Data type: statistical data
Explanation note: Maximum likelihood bootstrap percentages (BS) are provided at each node. Thickened lines indicate Bayesian inference posterior probability (PP) ≥ 0.9.
Figure
Data type: statistical data
Explanation note: Maximum likelihood phylogram of Polypodiineae obtained from the rps4-trnS dataset. Maximum likelihood bootstrap percentages (BS) are provided at each node. Thickened lines indicate Bayesian inference posterior probability (PP) ≥ 0.9.
Figure
Data type: statistical data
Explanation note: Maximum likelihood bootstrap percentages (BS) are provided at each node. Thickened lines indicate Bayesian inference posterior probability (PP) ≥ 0.9.