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Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae.

Author information

Affiliations

  • 1. Division of Silviculture, Taiwan Forestry Research Institute, 53 Nan-Hai Rd., Taipei 100, Taiwan.
      Authors
    • Chen CW 1
    • Huang YM 1
    (2 authors)
  • 2. The Pringle Herbarium, Department of Plant Biology, The University of Vermont, 27 Colchester Ave., Burlington, VT 05405, USA.
      Authors
    • Sundue M 2
    (1 author)
  • 3. Institute of Ecology and Evolutionary Biology, National Taiwan University, No. 1, Sec. 4, Roosevelt Road, Taipei, 10617, Taiwan.
      Authors
    • Kuo LY 3
    (1 author)
  • 4. Natural photographer, 664, Hu-Shan Rd., Caotun Township, Nantou 54265, Taiwan.
      Authors
    • Teng WC 4
    (1 author)

Phytokeys, 07 Apr 2017, (78):83-107
https://doi.org/10.3897/phytokeys.78.12040 PMID: 28781553 PMCID: PMC5543276

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Abstract 

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.

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Logo of phytoPensoft PublishersAboutEditorial TeamAuthor GuidelinesSubmissionPhytoKeys
PhytoKeys. 2017; (78): 83–107.
Published online 2017 Apr 7. https://doi.org/10.3897/phytokeys.78.12040
PMCID: PMC5543276
PMID: 28781553

Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae

Cheng-Wei Chen,1 Michael Sundue,2 Li-Yaung Kuo,3 Wei-Chih Teng,4 and Yao-Moan Huang1
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Abstract

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.

Keywords: Fern, morphology, Papua New Guinea, phylogeny, recircumscription, taxonomy, the Solomon Islands

Introduction

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 (Copeland 1947; Fig. ​Fig.1A).1A). Christensen (1937) was the first to point out that D. phaeostigma had been independently described under three different genera or subgenera. All told, generic placements has included Aspidium (≡ Tectaria) (Cesati 1877, Baker 1891), Dryopteris (Christensen 1906, Alderwerelt van Rosenburgh 1908, Copeland 1911, Brause 1920, Alderwerelt van Rosenburgh 1924), and Polystichum (Rosenstock 1911). Copeland (1947) inaugurated the new monotypic genus Dryopolystichum in his Genera Filicum, and argued that it was closest to Ctenitis. Pichi Sermolli (1977) agreed, citing the ctenitoid rachis, free venation, and peltate indusium as critical characters. Holttum included the genus in his “Tectarioid Group” in his list of Malaysian pteridophytes (Holttum 1959), but then omitted it in his 1991 treatment of that group.

An external file that holds a picture, illustration, etc. Object name is phytokeys-78-083-g001.jpg

Dryopolystichum phaeostigma (based on SITW10443). A Habitat B Plants C Peltate indusia D Venation E Sulcate rachis-costa architecture F Longitudinal section of the rhizome.

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 (Little and Barrington 2003). A peltate indusium is diagnostic of polystichoid ferns, including Phanerophlebia and Polystichum, but also found in a few distantly related genera in Polypodiineae such as Cyclodium, Cyclopeltis, Rumohra, Megalastrum, and Tectaria (Kramer and Green 1990).

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) (Sundue and Rothfels 2014, PPG I 2016). However, the remaining prominent features including pinnate-pinnatifid leaf dissection (Fig. ​(Fig.1B),1B), peltate indusium (Fig. ​(Fig.1C),1C), catadromous free veins (Fig. ​(Fig.1D),1D), and sulcate rachis-costa architecture (Fig. ​(Fig.1E),1E), do not clearly place it within any Polypodiinae family (Christensen 1937, Copeland 1947).

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. ​(Fig.1D).1D). This character is relatively uncommon in the Polypodiineae. It can be found in Dryopteridaceae, mostly in Megalastrum, and less commonly in Stigmatopteris, Ctenitis, and Pleocnemia (Moran et al. 2014, Moran and Labiak 2016). It can also be found in some Tectariaceae such as Pteridrys and Tectaria (Ding et al. 2014). Among these genera, Pleocnemia seems morphologically the most similar to Dryopolystichum because its rachises are adaxially sulcate and narrowly winged laterally. Pleocnemia, however, lacks a peltate indusium (Holttum 1974).

Subsequent to its establishment as a new genus in Genera Filicum (Copeland 1947), and Sermolli’s (1977) contribution, no other substantial argument was made for generic placement of Dryopolystichum. More recent studies maintained Dryopolystichum as a distinct genus, placing it under Dryopteridaceae (Kramer and Green 1990, Smith et al. 2006, Christenhusz et al. 2011). The recently published community-derived classification for extant lycophytes and ferns also places Dryopolystichum in the Dryopteridaceae but without assigning it to subfamily (PPG I 2016).

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.

Materials and methods

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 Chen et al. (2014).

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 (Kuo et al. 2017). The genome size of spore nuclei should be half the genome size of leaf nuclei in the case of sexual and the same size in the case of apomictic reproduction (Kuo et al. 2017). We followed Kuo et al. (2017) for the extraction of leaf nuclei. For extraction of spore nuclei, we used an optimized bead-vortexing treatment with vertex duration of 1 minute and vertex speed of 1,900 rpm, as described by Kuo et al. (2017). An external standard was not necessary since we only need to compare the two phases of the life-cycle to each other.

DNA extraction, amplification and sequencing

Total DNA was extracted using a modified CTAB-Qiagen column protocol (Kuo 2015). Three plastid DNA regions, rbcL, rps4-trnS (rps4 gene + rps4-trnS intergenic spacer), and trnL-F (trnL gene + trnL-trnF intergenic spacer), were amplified and sequenced using the primers “ESRBCL1F” and “1379R” for rbcL (Pryer et al. 2001, Schuettpelz and Pryer 2007), “RPS5F” and “TRNSR” for rps4-trnS (Nadot et al. 1995, Smith and Cranfill 2002), and “FernL 1Ir1” and “f” for trnL-F (Taberlet et al. 1991, Li et al. 2010).

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.

DNA alignment and phylogenetic analyses

Initial BLAST against the NCBI nucleotide database (Altschul et al. 1990) based on rbcL sequences indicated that Dryopolystichum phaeostigma is closely related to the species of Polypodiineae families, including Lomariopsidaceae, Nephrolepidaceae, Tectariaceae, and Dryopteridaceae. Accordingly, we assembled a data matrix including 250 species representing 36 genera from these families (Appendix). Sampling included all the four genera in which D. phaeostigma has been placed (i.e., Dryopteris, Polystichum, and Tectaria).

Sequences were aligned using Geneious v6.1.8 (Drummond et al. 2011) and then manually checked for errors. The three single-region (rbcL, rps4-trnS, and trnL-F) and dataset combining all three were independently subjected to both maximum likelihood (ML) and Bayesian inference (BI) phylogenetic analyses. Data matrices are available in TreeBASE, study number 20506, at https://treebase.org/. ML tree searches were conducted using RAxML (Stamatakis 2006) employing the GTRGAMMA substitution model through the CIPRES portal (Miller et al. 2010). Five independent searches for the ‘best tree’ and 1,000 bootstrap replicates were performed using a region-partitioned dataset. BI analyses were conducted using MrBayes 3.2.1 (Ronquist and Huelsenbeck, 2003) employing the same substitution model as in ML analysis. Each analysis consisted of two independent runs with four chains for 106 generations, sampling one tree every 1000 generations. Burn-in was set to 10000 based on our preliminary analysis. The convergences of MCMC runs were checked using Tracer v.1.6 (Rambaut et al. 2014).

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 (Philippe et al. 2005), we analyzed the concatenated dataset under MP in order to compare those results with our ML phylogeny. The MP analyses were conducted using TNT (Goloboff et al. 2008) following the search strategy detailed in Sundue et al. (2014).

Results

Phylogenetic analyses

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 2, 3). In comparison, the trnL-F phylogeny placed D. phaeostigma sister to Cyclopeltis (BS = 74%), and Dryopolystichum + Cyclopeltis was sister to Dracoglossum + Lomariopsis (Suppl. material 4). There was no strongly supported conflict between the ML and BI phylogenies (Suppl. materials 1–4). Both the ML and BI phylogenies based on the combined dataset (Fig. ​(Fig.2,2, Suppl. material 1) reveal the same topology as those based on the rbcL and rps4-trnS regions. Bootstrap support and posteriori probability (PP) for the above relationships were generally very high except for the branches placing D. phaeostigma, where BS was ≤ 70% and PP were ≤ 0.9 in all the phylogenies.

An external file that holds a picture, illustration, etc. Object name is phytokeys-78-083-g002.jpg

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 1. Voucher information and GenBank accession numbers are shown in Appendix.

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).

Karyology, reproductive mode, and spore measurements

All examined sporangia (SITW10443) produced 32 normal spores, and the mean spore length was 64.1 ± 4.5 μm (Fig. ​(Fig.3).3). The chromosome number of the three sporophyte somatic cells observed was ca. 164 (Fig. ​(Fig.4).4). Results of flow cytometry revealed that the genome size of spore nuclei is approximately half of those of leaf nuclei (Fig. ​(Fig.55).

An external file that holds a picture, illustration, etc. Object name is phytokeys-78-083-g003.jpg

Spores SEM of Dryopolystichum phaeostigma. A Lateral view of the spore B Detail of surface. Scale bars: A = 50 μm, B = 10 μm.

An external file that holds a picture, illustration, etc. Object name is phytokeys-78-083-g004.jpg

Chromosome number of Dryopolystichum phaeostigma. A Chromosomes at mitosis metaphase, 2n = ca. 164 (SITW10443) B explanatory illustration of A. Scale bars = 10 μm.

An external file that holds a picture, illustration, etc. Object name is phytokeys-78-083-g005.jpg

Relative DNA contents of Dryopolystichum phaeostigma spore and leaf nuclei inferred by flow cytometry.

Discussion

Phylogenetic placement of Dryopolystichum

The reconstructed maximum likelihood and Bayesian inference phylogenies unambiguously resolved Dryopolystichum within Lomariopsidaceae (Fig. ​(Fig.2),2), a position not previously suggested (Kramer and Green 1990, Smith et al. 2006, Christenhusz et al. 2011, PPG I 2016). This placement is consistent in all our analyses. Nonetheless, the generic position of Dryopolystichum within Lomariopsidaceae remains poorly resolved. This uncertainty may be partially explained by the incongruence between trnL-F and the other analyzed regions, but our process of removing the long-branched genera showed that low BS was retrieved only when Dryopolystichum and Lomariopsis were both included in the analysis. These results may also be explained by the large amounts of missing data in Lomariopsis; 19 of the 25 species included were represented by trnL-F data alone. We recommend further phylogenetic study using an expanded dataset to resolve the intergeneric relationships within Lomariopsidaceae.

Recircumscription of Lomariopsidaceae

Phylogenetic analyses using DNA sequences have served as the basis for redrawing fern classifications in the 21th century (Smith et al. 2006, Christenhusz et al. 2011, PPG I 2016). With respect to family circumscription, one of the most dramatically changed families is Lomariopsidaceae (Tsutsumi and Kato 2006, Schuettpelz and Pryer 2007, Christenhusz et al. 2013). Just prior to the molecular era, Lomariopsidaceae was treated as one of the largest fern families with six genera and over 500 species (e.g., Kramer and Green 1990) and was strongly supported by the following combination of characters: rhizomes with ventral root insertion, dictyosteles with elongate ventral meristeles, and dimorphic leaves where the fertile leaves had acrostichoid sori (Holttum and Hennipman 1959, Kramer and Green 1990).

Subsequent molecular phylogenetic analyses demonstrated that most genera previously treated in Lomariopsidaceae should be transferred to Dryopteridaceae (Tsutsumi and Kato 2006, Schuettpelz and Pryer 2007). The combination of characters uniting the former Lomariopsidaceae are now interpreted to have evolved multiple times, and to be correlated with dorsiventrality of the rhizome (Moran et al. 2010, McKeown et al. 2012). Meanwhile, Cyclopeltis was transferred from Dryopteridaceae to Lomariopsidaceae as suggested by molecular phylogeny (Schuettpelz and Pryer 2007), although it has none of the characters formerly used to circumscribe Lomariopsidaceae (Holttum and Hennipman 1959, Kramer and Green 1990).

More recently, the neotropical genus Dracoglossum was established (Christenhusz 2007) and later transferred to Lomariopsidaceae from Tectariaceae based on a molecular phylogeny (Christenhusz et al. 2013). This pattern was also unexpected since there are essentially no shared morphological characters by Dracoglossum and Lomariopsis, except for the ribbon-like gametophyte (R. C. Moran pers. com.). Our finding, that Dryopolystichum belongs to Lomariopsidaceae, comes as a further surprise. With these changes, Lomariopsidaceae is a family of five genera (Cyclopeltis, Dracoglossum, Dryopolystichum, Lomariopsis, and Thysanosoria) and ca. 70 species. As far as we can tell, none of the morphological traits commonly used unify these genera (Table ​(Table1).1). In the following paragraphs, we provide a recircumscription of both Lomariopsidaceae and Dryopolystichum, and then discuss selected characters in the light of our phylogenetic placement.

Table 1.

Comparison of morphological characters of the five Lomariopsidaceae genera [based on Holttum and Hennipman (1959), Holttum (1991), Roubik and Moreno (1991), Moran (2000), Christenhusz (2007), Rouhan et al. (2007), and this study].

Genera Cyclopeltis Dracoglossum Dryopolystichum Lomariopsis Thysanosoria
Habitterrestrialterrestrialterrestrialhemiepiphytehemiepiphyte
Rhizomeerectshort creepingerectclimbingclimbing
Frond division*pinnatesimplepinnate-pinnatifidpinnatepinnate
Pinnae articulationarticulate–not articulatearticulatearticulate
Venationfreereticulate, with included veinletfreefreefree
Rachis-costa architectureprominentprominentgroovedgrooved or flatgrooved
Sporangiaform rounded soriform rounded soriform rounded soriacrostichoidform rounded sori
Indusiapeltate if presentpeltate if presentpeltateabsentabsent
Perine ornamentationbroad foldsnarrow crestsnarrow crestsvariousbroad folds

*matured plant, -not applicable

Taxonomic treatment

Lomariopsidaceae

Keywords: Plantae, ORDO, FAMILIA

Alston, Taxon 5(2): 25. 1956.

Type.

Lomariopsis Fée, Mém. Foug., 2. Hist. Acrostich.: 10. 1845.

Description.

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 (Holttum and Hennipman 1959), but it has not been, to the present, subject to molecular phylogenetic analysis.

Dryopolystichum

Keywords: Plantae, ORDO, FAMILIA

Copel., Gen. Fil. 125, t. 4. 1947.

Type.

Dryopolystichum phaeostigma (Ces.) Copel., Gen. Fil. 125, t. 4. 1947.

Description.

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.

Dryopolystichum phaeostigma

Keywords: Plantae, ORDO, FAMILIA

(Ces.) Copel., Gen. Fil. 125, t. 4. 1947.

  • 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]).

Type.

Based on Aspidium phaeostigma Ces.

Description.

Equal to the genus.

Distribution.

New Guinea, the Bismark archipelago, and the Solomon Islands.

Comparison of selected characters of Dryopolystichum

Perine architecture of Dryopolystichum is very similar to that of Dracoglossum plantagineum (Christenhusz 2007, Fig. ​Fig.3).3). They are loosely attached, forming thin crests, and having a spiculate microstructure. Perine of Cyclopeltis and Thysanosoria are also similar in being loosely attached and having a spiculate microstructure, but they differ by having broader folds (Holttum and Hennipman 1959, Tryon and Lugardon 1991). The perine characters, however, are not shared by all the taxa of Lomariopsidaceae especially considering the variation of ornamentation existing in Lomariopsis (Rouhan et al. 2007). Moreover, these perine characters also appear in other Polypodiineae lineages particularly in bolbitidoid ferns (Moran et al. 2010) as well as in various Aspleniineae lineages (Sundue and Rothfels 2014, PPG I 2016).

Blackish sclerenchyma strands are visible in the rhizome sections of Dryopolystichum (Fig. ​(Fig.1F).1F). These are also present in Dracoglossum, Cyclopeltis, and Lomariopsis, but similar characters are known from various groups throughout Polypodiineae (Hennipman 1977, Moran 1986, Hovenkamp 1998). Further studies might reveal variation in these strands to be of systematic value.

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 (Holttum and Hennipman 1959, Moran 2000). In contrast, Dracoglossum and Cyclopeltis have non-winged and non-sulcate rachises (Holttum 1991, Christenhusz 2007).

The chromosome number in somatic cells of Dryopolystichum phaeostigma was ca. 164 (Fig. ​(Fig.4).4). The base numbers for Lomariopsidaceae genera (Cyclopeltis, Dracoglossum, and Lomariopsis) are 40 or 41 (Walker 1985, Kato and Nakato 1999, Moran 2000), suggesting that D. phaeostigma is a tetraploid.

Our flow cytometry and spore count results indicate that Dryopolystichum phaeostigma is sexually reproducing and has 32 spores per sporangium (Fig. ​(Fig.5).5). In Polypodiales, sporogenesis leading to the formation of 64 spores in a sporangium is by far the most common pattern of sexually reproducing species, e.g., Aspleniaceae (Gabancho et al. 2010), Athyriaceae (Kato et al. 1992, Takamiya et al. 1999), Davalliaceae (Chen et al. 2014), Dryopteridaceae (Lu et al. 2006), Polypodiaceae (Wang et al. 2011), Pteridaceae (Huang et al. 2006), and Thelypteridaceae (Ebihara et al. 2014). Cases of sporogenesis resulting in 32 spores per sporangium are known from a few Polypodiales ferns but all belong to the suborders Lindsaeineae and Pteridineae, i.e., Lindsaeaceae (Lin et al. 1990), Cystodiaceae (Gastony 1981), and Ceratopteris (Pteridaceae; Lloyd 1973). Our study provides the first confirmed case of a sexual reproduction with 32 spores per sporangium in the suborder Polypodiineae.

Conclusion

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.

Supplementary Material

XML Treatment for Lomariopsidaceae :
XML Treatment for Dryopolystichum :
XML Treatment for Dryopolystichum phaeostigma :

Acknowledgements

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.

Appendix

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.

TaxonGenbank accession numbers
rbcL rps4-trnS trnL-F
Dryopteridaceae
Arachniodes aristata (G.Forst.) TindaleKJ464418–KJ464592
Arachniodes denticulata (Sw.) ChingKJ464419–KJ464593
Arthrobotrya articulata J.Sm.–GU376714GU376565
Arthrobotrya wilkesiana Copel.–GU376719GU376569
Bolbitis acrostichoides (Afzel.) ChingKJ464420GU376644GU376500
Bolbitis aliena (Sw.) Alston–GU376646GU376502
Bolbitis angustipinna (Hayata) H.Ito–GU376654GU376509
Bolbitis appendiculata (Willd.) K.Iwats.–GU376647GU376503
Bolbitis auriculata (Lam.) AlstonKJ464421GU376649GU376505
Bolbitis bipinnatifida (J.Sm.) K.Iwats.–GU376676GU376530
Bolbitis fluviatilis (Hook.) Ching–GU376656GU376510
Bolbitis gemmifera (Hieron.) C.Chr.–GU376657GU376511
Bolbitis heteroclita (Pr.) Ching–GU376659GU376513
Bolbitis heudelotii (Bory) Alston–GU376662GU376515
Bolbitis humblotii (Baker) ChingKJ464422GU376663GU376516
Bolbitis lonchophora (Kunze) C.Chr.–GU376664GU376517
Bolbitis major (Bedd.) Hennipman–GU376665GU376518
Bolbitis portoricensis (Sprengel) Hennipman–GU376670GU376523
Bolbitis salicina (Hook.) Ching–GU376671GU376525
Bolbitis semipinnatifida (Fée) Alston–GU376672GU376526
Bolbitis serratifolia (Mertens) Schott–GU376673GU376527
Bolbitis sinuata (C.Presl) Hennipman–GU376675GU376529
Bolbitis tibetica Ching & S.K.Wu–GU376677GU376531
Ctenitis eatonii (Baker) ChingKF709483–KJ196645
Ctenitis sinii (Ching) Ohwi––KJ196643
Ctenitis subglandulosa (Hance) Ching––KJ196655
Ctenitis yunnanensis Ching & Chu H.Wang––KJ196715
Cyclodium heterodon (Schrad.) T. Moore var. heterodonKJ464425–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–GU376678KJ196632
Dryopteris wallichiana (Spreng.) Hyl.KJ464428GU376680KJ464599
Elaphoglossum amygdalifolium (Mett.) Christ–GU376681–
Elaphoglossum burchellii (Baker) C.Chr.–GU376682GU376533
Elaphoglossum decoratum (Kunze) T.MooreKJ464429GU376683KJ464600
Elaphoglossum guentheri Rosenst.–GU376684GU376535
Elaphoglossum langsdorffii T.Moore–GU376685GU376536
Elaphoglossum lloense (Hook.) T.Moore–GU376686GU376537
Elaphoglossum luridum Christ––GU376538
Elaphoglossum squamipes (Hook.) T.Moore––GU376539
Lastreopsis amplissima (C.Presl) TindaleKJ464432–KJ464604
Lastreopsis decomposita (R.Br.) TindaleKJ464439––
Lastreopsis hispida (Sw.) TindaleKJ464446–KJ464614
Lastreopsis killipii (C.Chr. & Maxon) TindaleKJ464448KF709505–
Lastreopsis marginans (F.Muell.) TindaleKJ464449GU376691KJ464616
Lastreopsis poecilophlebia (Hook.) Labiak, Sundue & R.C.MoranKJ464423GU376692KJ464594
Lastreopsis tenera (R.Br.) TindaleKJ464467GU376699KJ464636
Lastreopsis tripinnata (F.Muell. ex Benth.) Labiak, Sundue & R.C.MoranKJ464491GU376700–
Lastreopsis walleri TindaleKJ464472GU376701–
Lastreopsis wurunuran (Domin) TindaleKJ464474GU376704–
Lomagramma brooksii Copel.–GU376705GU376542
Lomagramma cordipinna Holttum–GU376707GU376543
Lomagramma lomarioides (Blume) J.Sm.––GU376550
Lomagramma matthewii (Ching) HolttumKJ464476–KJ464640
Lomagramma perakensis Bedd.––GU376552
Lomagramma pteroides J.Sm.––GU376555
Lomagramma sinuata C.Chr.––GU376556
Lomagramma sumatrana Alderw.––GU376558
Maxonia apiifolia (Sw.) C.Chr.KJ464477GU376709KJ464641
Megalastrum abundans (Rosenst.) A.R.Sm. & R.C.MoranKJ464478–KJ464642
Megalastrum atrogriseum (C.Chr.) A.R.Sm. & R.C.MoranKJ464479GU376710KJ464643
Megalastrum connexum (Kaulf.) A.R.Sm. & R.C.MoranKJ464481–KJ464645
Megalastrum lanatum (Fée) HolttumKJ464483–KJ464647
Megalastrum littorale R.C.Moran, J.Prado & Labiak–GU376651GU376561
Megalastrum macrotheca (Fée) A.R.Sm. & R.C.MoranKJ464484GU376697KJ464648
Megalastrum vastum (Kunze) A.R.Sm. & R.C.MoranKJ464487GU376658KJ464651
Mickelia bernoullii (Kuhn ex Christ) R.C.Moran, Labiak & Sundue–GU376666GU376506
Mickelia guianensis (Aubl.) R.C.Moran, Labiak & Sundue–GU376667GU376548
Mickelia hemiotis (Maxon) R.C.Moran, Labiak & Sundue––GU376512
Mickelia nicotianifolia (Sw.) R.C.Moran, Labiak & Sundue–KF667557GU376519
Mickelia oligarchica (Baker) R.C.Moran, Labiak & SundueKJ464489–GU376520
Mickelia scandens (Raddi) R.C. Moran, Labiak & Sundue–GU376696GU376547
Olfersia cervina KunzeKJ464493DQ153079KJ464652
Parapolystichum acuminatum (Houlston) Labiak, Sundue & R.C.MoranKJ464430KC977454KJ464601
Parapolystichum boivinii (Baker) RouhanKJ464435–KJ464607
Parapolystichum confine (Maxon ex C.Chr.) Labiak, Sundue & R.C.MoranKJ464438––
Parapolystichum effusum (Sw.) ChingKJ464441––
Parapolystichum effusum (Sw.) Ching subsp. divergens (Willd. ex Schkuhr) TindaleKJ464440––
Parapolystichum excultum (Mett.) Labiak, Sundue & R.C.Moran–KF709501GU376541
Parapolystichum glabellum (A.Cunn.) Labiak, Sundue & R.C.MoranKJ464445KF709503KJ464613
Parapolystichum microsorum (Endl.) Labiak, Sundue & R.C.MoranKJ464451GU376712KJ464617
Parapolystichum perrierianum (C.Chr.) RouhanKJ464455–KJ464623
Parapolystichum rufescens (Blume) Labiak, Sundue & R.C.MoranKJ464461–KJ464629
Parapolystichum vogelii (Hook.) RouhanKJ464470
Parapolystichum windsorensis (D.L.Jones & B.Gray) Labiak, Sundue & R.C.MoranKJ464473–KJ464639
Pleocnemia conjugata C.Presl–GU376713KF709510
Pleocnemia cumingiana C.PreslKJ196828–KJ196705
Pleocnemia dahlii (Hieron.) HolttumKJ196829–KJ196706
Pleocnemia hemiteliiformis (Racib.) HolttumKF709482KF667560KF709511
Pleocnemia irregularis (C.Presl) HolttumKF709491–KF709513
Pleocnemia leuzeana (Gaudich.) C.PreslKJ196830––
Pleocnemia olivacea (Copel.) HolttumKJ464495––
Pleocnemia presliana HolttumKJ464496KF667561–
Pleocnemia rufinervis NakaiJF303976KF667562–
Pleocnemia winitii HolttumEF460686–KF709515
Polybotrya alfredii BradeKJ464497KF667563KJ464653
Polybotrya andina C.Chr.KJ464498KP271084KJ464654
Polybotrya pubens Mart.KJ464499KP271085–
Polystichum tsus-simense (Hook.) J.Sm. var. mayebarae (Tagawa) Sa.KurataAB575224–DQ150408
Pseudotectaria biformis (Mett.) Holttum––KF897951
Pseudotectaria decaryana (C.Chr.) Tardieu––KF897952
Rumohra adiantiformis (G.Forst.) ChingKJ464500–KJ464655
Rumohra berteroana (Colla) J.J. Rodr.KJ464503–KJ464657
Stigmatopteris ichthiosma (Sodiro) C.Chr.KJ464504–KJ464658
Stigmatopteris killipiana LellingerKJ464505–KJ464659
Stigmatopteris lechleri (Mett) C.Chr.KJ464506KP271087KJ464660
Stigmatopteris sordida (Maxon) C.Chr.KJ464507–KJ464661
Teratophyllum koordersii Holttum––GU376566
Teratophyllum ludens (Fée) Holttum––GU376567
Teratophyllum wilkesianum HolttumKJ464508––
Nephrolepidaceae
Nephrolepis abrupta (Bory) Mett.HM748137KF667559–
Nephrolepis acutifolia (Desv.) Christ.HM748139––
Nephrolepis biserrata (Sw.) SchottAB575227GU376688–
Nephrolepis brownii (Desv.) Hovenkamp & Miyam.KR816691––
Nephrolepis cordifolia (L.) C.PreslAB575228––
Nephrolepis davalliae Alderw.HM748147––
Nephrolepis davallioides KunzeHM748148GU376690–
Nephrolepis exaltata (L.) SchottHM748149––
Nephrolepis falcata (Cav.) C.Chr.HM748150––
Nephrolepis falciformis J.Sm.AB232404––
Nephrolepis lauterbachii (Christ) ChristHM748153––
Nephrolepis pectinata (Willd.) SchottHM748155––
Nephrolepis pendula (Raddi) J.Sm.HM748156––
Nephrolepis radicans (Burm.) KuhnHM748157––
Nephrolepis rivularis (Vahl) Mett.HM748158––
Nephrolepis undulata J.Sm.HM748159––
Lomariopsidaceae
Cyclopeltis crenata (Fée) C.Chr.DQ054517EF540718DQ51448
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–KJ628952DQ396560
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.) KuhnAY818677–DQ396578
Lomariopsis maxonii (Underw.) Holttum––DQ396580
Lomariopsis muriculata Holttum––DQ396582
Lomariopsis palustris (Hook.) Mett. ex Kuhn–HM748162DQ396585
Lomariopsis pervillei Kuhn––DQ396586
Lomariopsis pollicina (Willemet) Mett. ex KuhnEF463235–DQ396588
Lomariopsis prieuriana Fée––DQ396590
Lomariopsis recurvata Fée––DQ396592
Lomariopsis rossii Holttum––DQ396594
Lomariopsis salicifolia (Kunze) Lellinger––DQ396595
Lomariopsis sorbifolia (L.) FéeEF463236––
Lomariopsis spectabilis Mett.AB232401–KJ196685
Lomariopsis vestita E.Fourn.––DQ396598
Lomariopsis wrightii Mett.––DQ396600
Tectariaceae
Arthropteris altescandens J.Sm.KF667636KF667550KF667606
Arthropteris articulata (Brack.) C.Chr.KC977367KC977437KC977411
Arthropteris beckleri (Hook.) Mett.U05605–KF667607
Arthropteris cameroonensis AlstonKF667638––
Arthropteris guinanensis H.G.Zhou & Y.Y.HuangKC977364KC977442KC977404
Arthropteris monocarpa (Cordem.) C.Chr.HM748132–KF897941
Arthropteris orientalis (Gmel.) Posth.HM748133KC977435KC977420
Arthropteris palisotii (Desv.) AlstonAB575230KC977427KC977406
Arthropteris parallela (Baker) C.Chr.EF463266KC977453KC977425
Arthropteris paucivenia (C.Chr.) H.M.Liu, Hovenkamp & H.Schneid.EF463268–KC977426
Arthropteris repens (Brack.) C.Chr.KC977368KC977438KC977412
Arthropteris tenella (G.Forst.) J.Sm. ex Hook.f.KC977363KF011547KC977424
Hypoderris brauniana (H.Karst.) F.G.Wang & Christenh.KF667647–KF667618
Hypoderris brownii J.Sm.KF667642–KF667611
Hypoderris nicotianifolia (Baker) R.C.Moran, Labiak & J.PradoKF667653–KF667626
Pteridrys australis ChingKJ196892–KJ196678
Pteridrys cnemidaria (Christ) C.Chr. & ChingKF709488–KF709517
Pteridrys lofouensis (Christ) C.Chr. & ChingEF460687KF667566–
Pteridrys microthecia (Fée) C.Chr. & ChingKJ196848–KF709518
Pteridrys syrmatica (Willd.) C.Chr. & ChingKJ196875–KF709519
Tectaria acerifolia R.C.MoranKF887170–KF897954
Tectaria angulata (Willd.) Copel.KJ196876–KJ196656
Tectaria aurita (Sw.) S.ChandraKJ196849–KJ196631
Tectaria barberi (Hook.) Copel.KJ196846–KJ196628
Tectaria borneensis S.Y.DongKJ196854KF667555KJ196642
Tectaria cicutaria (L.) Copel.KF667649–KF667620
Tectaria coadunata (J.Sm.) C.Chr.KJ196851–KJ196661
Tectaria crenata Cav.KF667650KF667568KF667621
Tectaria decurrens (C.Presl) Copel.AB575232–DQ514524
Tectaria devexa (Kunze ex Mett.) Copel.AB575233KP271088KF897956
Tectaria dilacerata (Kunze) MaxonKF887173–KF897957
Tectaria fauriei TagawaAB575234–KJ196658
Tectaria fernandensis C.Chr.KF887174–KF897958
Tectaria gigantea (Blume) Copel.KJ196853–KJ196660
Tectaria griffithii (Baker) ChingKF667652–KF667624
Tectaria grossedentata Ching & Chu H.WangKJ196882KP271089KJ196667
Tectaria harlandii (Hook.) C.M.KuoAB575231–KJ196648
Tectaria harlandii (Hook.) C.M.KuoKF887178–KF897961
Tectaria heracleifolia (Willd.) Underw.KF887180–KF897963
Tectaria herpetocaulos Ching & Chu H. WangKJ196884–KJ196669
Tectaria heterocarpa C.V.MortonKF887181–KF897964
Tectaria impressa (Fée) HolttumKJ196841–KF897965
Tectaria kusukusensis (Hayata) LellingerEF460681–KF897968
Tectaria labrusca (Hook.) Copel.KJ196818–KJ196692
Tectaria luchunensis S.K.WuKJ196845KP271090KJ196627
Tectaria macleanii (Copel.) S.Y.DongKJ196810–KJ196680
Tectaria melanocaula (Blume) Copel.KJ196832–KJ196709
Tectaria morsei (Baker) P.J.Edwards ex S.Y.DongKJ196893KF667570KF561675
Tectaria nayarii MazumdarEF463267–KJ196699
Tectaria paradoxa (Fée) SledgeKF887189–KF897971
Tectaria phaeocaulis (Rosenst.) C.Chr.AB232397KF709499KF897972
Tectaria pica (L.) C.Chr.KF887191GU376715KF897973
Tectaria polymorpha (Wall. ex Hook.) Copel.KJ196888GU376716KJ196657
Tectaria prolifera (Hook.) R.M.Tryon & A.F.TryonEF463273–KF897974
Tectaria psomiocarpa S.Y.DongKJ196822KF667572KJ196698
Tectaria pubens R.C.MoranKF887193KF667573KF897975
Tectaria quinquefida (Baker) ChingKJ196885–KJ396622
Tectaria repanda (Willd.) HolttumKJ196831–KJ196707
Tectaria sagenioides (Mett.) Christenh.KF887194KF667575KF561672
Tectaria semipinnata (Roxb.) MortonKJ196817KF667577KJ196691
Tectaria simonsii (Baker) ChingAB575236–KF897977
Tectaria singaporiana (Wall. ex Hook. & Grev.) ChingKF887196–KF897978
Tectaria subglabra (Holttum) S.Y.Dong––KJ196676
Tectaria subsageniacea (Christ) Christenh.KF887197KF667576KF561670
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.WalkerKF887201–KF897983
Tectaria zeilanica (Houtt.) SledgeAB232395–KF709521
Triplophyllum crassifolium HolttumKF887203–KF897985
Triplophyllum fraternum (Mett.) HolttumKF667657–KF667630
Triplophyllum funestum (Kunze) HolttumEF463276–KF667631
Triplophyllum glabrum J.Prado & R.C.MoranKF887207–KF897989
Triplophyllum heudelotii Pic.Serm.––KF897990
Triplophyllum jenseniae (C.Chr.) HolttumKF667660–KF667633
Triplophyllum pentagonum (Bonap.) HolttumKF667662–KF667635
Triplophyllum pilosissimum (J.Sm. ex T.Moore) Holttum––KU605127
Triplophyllum securidiforme (Hook.) Holttum––KU605128
Triplophyllum vogelii (Hook.) HolttumKF667661–KF667634
Oleandraceae
Oleandra articulata (Sw.) C.PreslKF667644KF709500KF667613
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 & TsutsumiJX103718KC914565–
Polypodiaceae
Campyloneurum minus FéeKF667665––
Microgramma lycopodioides (L.) Copel.KF667664––
Niphidium longifolium (Cav.) C.V.Morton & LellingerKF667663KF709495–

Notes

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

Supplementary materials

Supplementary material 1

Figure S1. Maximum likelihood phylogram of Polypodiineae obtained from the combined (rbcL + rps4-trnS + trnL-F) dataset.

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.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Cheng-Wei Chen, Michael Sundue, Li-Yaung Kuo, Wei-Chih Teng, Yao-Moan Huang

Supplementary material 2

Figure S2. Maximum likelihood phylogram of Polypodiineae obtained from the rbcL dataset.

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.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Cheng-Wei Chen, Michael Sundue, Li-Yaung Kuo, Wei-Chih Teng, Yao-Moan Huang

Supplementary material 3

Figure S3. Maximum likelihood phylogram of Polypodiineae obtained from the rps4-trnS dataset.

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.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Cheng-Wei Chen, Michael Sundue, Li-Yaung Kuo, Wei-Chih Teng, Yao-Moan Huang

Supplementary material 4

Figure S4. Maximum likelihood phylogram of Polypodiineae obtained from the trnL-F dataset.

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.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Cheng-Wei Chen, Michael Sundue, Li-Yaung Kuo, Wei-Chih Teng, Yao-Moan Huang

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