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Research Article
The systematic position of Dryopteris blanfordii subsp. nigrosquamosa (Ching) Fraser-Jenkins within the genus Dryopteris Adans.
expand article infoAnastasiya A. Krinitsina‡, Maxim S. Belenikin§, Olga A. Churikova‡, Sergey V. Kuptsov‡, Maxim I. Antipin‡, Maria D. Logacheva‡, Anna S. Speranskaya‡
‡ Lomonosov Moscow State University, Moscow, Russia
§ Moscow Institute of Physics and Technology, Dolgoprudny, Russia

Corresponding author: Anastasiya A. Krinitsina ( ankrina@gmail.com )

Academic editor: Thais Almeida
© 2017 Anastasiya A. Krinitsina, Maxim S. Belenikin, Olga A. Churikova, Sergey V. Kuptsov, Maxim I. Antipin, Maria D. Logacheva, Anna S. Speranskaya.
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: Krinitsina AA, Belenikin MS, Churikova OA, Kuptsov SV, Antipin MI, Logacheva MD, Speranskaya AS (2017) Systematic position of Dryopteris blanfordii subsp. nigrosquamosa (Ching) Fraser-Jenkins within the genus Dryopteris Adans.. PhytoKeys 90: 89-112. https://doi.org/10.3897/phytokeys.88.14745
Open Access

Abstract

Dryopteris blanfordii (C.Hope) C.Chr. is a member of the Dryopteridaceae, growing in high altitude Picea or Abies forests (2900–3500 m) in China and India. Phylogenetic relationships between D. blanfordii subsp. nigrosquamosa and closely related species of Dryopteris were investigated using a combined analysis of multiple molecular data sets (the protein-coding region of rbcL and matK genes and intergenic spacers psbA-trnH, trnP-petG, rps4-trnS, trnL-trnF and rbcL-accD). An assumption about the position of D. blanfordii subsp. nigrosquamosa within Dryopteris was made by using the Maximum Likelihood and Bayesian Inference approach and chloroplast marker sequences of Dryopteris species from GenBank. The results demonstrated that Asian taxa D. blanfordii subsp. nigrosquamosa and D. laeta as well as two American species D. arguta and D. marginalis belong to the same clade, all four of them being part of Dryopteris section Dryopteris.

Keywords

chloroplast genome markers, Dryopteridaceae , Dryopteris blanfordii subsp. nigrosquamosa , phylogeny

Introduction

Dryopteris is a large fern genus of some 225–230 species belonging to the Dryopteridaceae (Fraser-Jenkins 1986, Kramer 1990, Zhang et al. 2012). Dryopteris blanfordii (C.Hope) C.Chr. grows in Picea or Abies forests at 2900–3500 m in China (Gansu, Sichuan, Xizang, Yunnan), Afghanistan, India, Kashmir, Nepal and Pakistan (Zhang et al. 2013). It is divided into two subspecies, nigrosquamosa (Ching) Fraser-Jenkins and blanfordii. Dryopteris blanfordii subsp. nigrosquamosa is abundant in India (the Kashmir valley) (Mir et al. 2014, Mir et al. 2015), China (Gansu, Sichuan, Xizang, Yunnan) and Nepal (Zhang et al. 2013).

The current taxonomy and infrageneric position of D. blanfordii are still unresolved. More than a quarter of a century ago, Fraser-Jenkins (1986) divided Dryopteris into several sections based on the comparison of macro- and micro-morphological traits. He established Sect. Remotae with a single species (D. blanfordii) in the Indian subcontinent and with two European species (Dryopteris remota Hayek. and D. corleyi Fraser-Jenk.). Fraser-Jenkins suggested that all species from Sect. Remotae are allopolyploids resulting from hybridization between species from different sections (such as sect. Fibrillosae, sect. Lophodium or sect. Marginatae) (Fraser-Jenkins 1986).

In a recent study, a phylogeny including 100 species of Dryopteris was reconstructed and 13 phylogenetic clades (or major evolutionary lineages) were identified using DNA sequences of four plastid loci (rbcL gene, rps4-trnS spacer, trnL intron and trnL-F spacer) (Zhang et al. 2012). Two of the three species, previously placed in Sect. Remotae, were identified in this study as members of other clades, namely clade Aemulae (D. corleyi) and clade Lophodium (D. remota) (Zhang et al. 2012). Similar results were demonstrated by Sessa et al. (2012a, 2012b), where seven plastid loci (rbcL, psbA-trnH, trnP-petG, rps4-trnS, trnL-F, trnG-trnR and rbcL-accD) and a single nuclear marker pgiC were used to analyse 97 Dryopteris species. The data of these phylogenies, although valuable, were far from complete and the taxonomic position of D. blanfordii subsp. nigrosquamosa remained unclear.

Material and methods

Plant material

To examine the morphology of D. blanfordii subsp. nigrosquamosa, adult plants from the Botanical Garden of Moscow State University were used. The parent plant of Dryopteris blanfordii (C.Hope) C.Chr. subsp. nigrosquamosa (Ching) Fraser-Jenk was collected in 2003 in Uttar Pradesh State, India, at 3000 m. Spores of the specimen were germinated under artificial conditions in the greenhouse complex of MSU Botanical Garden. Subsequently, developed sporophytes were transplanted to the outdoor section of the Botanical Garden. The adult specimens were used for DNA sampling. The voucher specimen was deposited at Herbarium MW. Reference morphological characters for D. blanfordii subsp. nigrosquamosa were scored from a type specimen (PE 00133945, locality: Tibet) and from descriptions of Dryopteris blanfordii (C.Hope) C.Chr. subsp. nigrosquamosa (Zhang et al. 2013, Mir et al. 2014, Mir et al. 2015).

Chloroplast markers sequencing and assembling

The chloroplast marker sequences of D. blanfordii subsp. nigrosquamosa were obtained during a large project on Polypodiales chloroplast genome sequencing. Sequencing data were generated using Illumina MiSeq high-throughput sequencing platform. For sample preparation, adult living plants were taken from the collection of the MSU Botanical Garden. The cpDNA fraction was extracted from 2.6 g (fresh weight) of fronds using a slightly modified cpDNA extraction protocol (Shi et al. 2012, Vieira et al. 2014). The purification of DNA was carried out using a protocol designed by the authors (Krinitsina et al. 2015). TruSeq protocol (NEBNext® DNA Library Prep Master Mix Set for Illumina, E6040, NEB reagents) was used for preparing the libraries. Pare end (PE) sequences (2×300bp) with a double number of each library reads about 1.2–1.97M were made. After quality trimming by Trimmomatic (Bolger et al. 2014), reads were filtered using 13 complete and five partial fern chloroplast genome sequences from RefSeq database and Bowtie2 (Langmead and Salzberg 2012). Then two sets of contigs were produced for both filtered and unfiltered sets of reads using Velvet Assembler (Zerbino and Birney 2008) and MIRA4 (Chevreux et al. 2004). Assembled contigs and scaffolds were used for assembling the complete chloroplast genome (the data are not presented in this paper) and for extracting target chloroplast markers, namely rbcL, matK genes and intergenic spacers psbA-trnH, trnP-petG, rps4-trnS, trnL-trnF and rbcL-accD.

Dryopteris blanfordii subsp. nigrosquamosa phylogenetic analysis

To determine the phylogenetic position of D. blanfordii subsp. nigrosquamosa, a phylogenetic analysis using sequences published in GenBank was performed. The GenBank accession numbers of sequences of Dryopteris species included in this study are listed in Appendix 1. Sequence alignment was conducted using Muscle algorithm and MEGA6.0 software package (www.megasoftware.net, Tamura et al. 2013). Phylogenetic analyses were performed using the Maximum Likelihood (ML) method at MEGA 6.0 (Tamura et al. 2013) and Bayesian Inference (BI) in BEAST (Bouckaert et al. 2014). A combined matrix including matK and rbcL gene and five intergenic spacers (psbA-trnH, rps4-trnS, trnL-trnF trnP-petG and rbcL-accD) of 84 Dryopteris species (including D. blanfordii subsp. nigrosquamosa) was analysed.

A bootstrapping of 1000 replicates for ML analysis was processed to estimate the confidence probabilities on each branch of the phylogenetic trees constructed. The initial tree (ML) for heuristic search was obtained by applying the Neighbour-Joining method to a matrix of pairwise distances estimated using the Maximum Composite Likelihood approach (Lindsay 1988, Tamura et al. 2013). All positions containing gaps and missing data were eliminated.

Bayesian analyses were run for 20,000,000 generations with four MCMC chains in two independent runs. The first 2,000,000 samples from each run were discarded as burn-in. Convergence was assessed by comparing the standard deviation of split frequencies between different runs (MCMC Trace Analysis Tool (Tracer) version v1.6.0 (Rambaut et al. 2014). For ML and BI analyses, optimal models of molecular evolution for combined matrices were identified using jModelTest2 (Darriba et al. 2012) through Bayesian Information Criterion (BIC).

Results

Dryopteris blanfordii subsp. nigrosquamosa phylogenetic position

Seven marker regions of the assembled cp genome were used for determining phylogenetic relationships between D. blanfordii subsp. nigrosquamosa and other Dryopteris species, i.e. protein-coding regions of rbcL and matK genes and intergenic spacers psbA-trnH, trnP-petG, rps4-trnS, trnL-trnF and rbcL-accD. These markers were assembled into a single data matrix consisting of 3734 total bases. The optimal model of molecular evolution for combined matrices was TPM1uf+G+I with BIC = 36592.7258. The phylogenetic tree is shown in Fig. 2. The analysis demonstrated close relationships between D. blanfordii subsp. nigrosquamosa, D. laeta, D. marginalis and D. arguta. The clades containing D. blanfordii subsp. nigrosquamosa were well-supported (≥80% bootstrap support). Dryopteris blanfordii subsp. nigrosquamosa is close to D. laeta (bootstrap=100/PP=100%), D. arguta (bootstrap=87/PP=99.6%) and D. marginalis (bootstrap=96/PP=100%). The results of Bayesian Inference analysis based on the combined matrix were highly congruent with the strict consensus tree from ML analysis. The clade that included D. arguta, D. marginalis, D. laeta and D. blanfordii subsp. nigrosquamosa in the combined matrix of seven markers had the posterior probability (PP) value of 100%.

The adult specimens of D. blanfordii analysed in the present work have 55–57×30–35cm fronds. The frond dissection is 2-pinnate with symmetrical pinnae and pinnules (Fig. 1A). The rachises and petioles are fibrillose and have dense basal scales. The scales on the petioles are dark-brown basally and light-brown at the apex (Fig. 1 B and C). The costa and rachises are slightly grooved adaxially (Fig. 1D) and rounded abaxially (Fig. 1E).

Figure 1. 

Dryopteris blanfordii subsp. nigrosquamosa morphology. A Frond of mature plants B Petiole covered with scales C Petiole scale D Adaxial surface of rachis and costa E Abaxial surface of rachis and costa.

Two closely related species, namely D. arguta and D. marginalis, are native to North America. Dryopteris marginalis is evergreen and has tawny or cinnamon-coloured scales, lanceolate and coriaceous laminae, with sori mostly at margins of ultimate pinnules’ segments (Table 1). Dryopteris arguta is winter green, having grassy-green to yellow-green, ovate-lanceolate, herbaceous, glandular laminae; the basal basiscopic pinnule and basal acroscopic pinnule are ± equal; its pinnule margins are serrate with spreading, spinelike teeth; the sori are medial. Both North American species have longer stipes (1/4–1/3 length of leaf), 1-pinnate-pinnatifid to 2-pinnate-pinnatifid fronds (Montgomery and Wagner 1993).

Table 1.
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Morphological characters of four species of Dryopteris: D. blanfordii subsp. nigrosquamosa, D. laeta, D. marginalis and D. arguta according to Montgomery and Wagner (1993), Mir et al. (2014), Mir et al. (2015).

Species D. blanfordii subsp. nigrosquamosa D. laeta D. marginalis D. arguta
Natural range Southeast Tibet, Western China, Nepal, India (Kashmir) North China, Eastern Siberia, North Korea and North Japan Eastern North America From British Columbia to Baja California
Seasonality semi-evergreen deciduous evergreen winter green
Rhizomes erect creeping ascending to erect short-creeping
Scale colour light brown with black veins pale brown tawny to cinnamon light brown
Lamina length (cm) 40–75 25–50 25–50 (75) 25–90
Lamina division 2-pinnate to 3-pinnate-pinnatifid 3-pinnate-pinnatifid 1-pinnate-pinnatifid to 2-pinnate-pinnatifid 2-pinnate-pinnatifid
Lamina colour and texture glaucous green, coriaceous green, herbaceous to thinly papyraceous green, coriaceous green to yellow-green, herbaceous, glandular
Lamina shape lanceolate to oblong-lanceolate ovate-oblong or deltoid-ovate ovate-lanceolate ovate-lanceolate
Stipe length 1/5-1/4 of rachis length 1/3 to 1/2 of rachis length 1/4 to 1/3 of rachis length 1/4 to 1/3 of rachis length
Sori arrangement in 1 row at each side of midvein, inframedial in 2 rows at each side of midvein in 1 row at each side of midvein, intramarginal at margins of segments in 1 row at each side of midvein, medial

Dryopteris laeta is characterised by a long stipe (length roughly equal to blade length) with very few lanceolate scales; deciduous, ovate-oblong or deltoid-ovate, 3-pinnate-pinnatifid, 25–50×15–40cm, herbaceous to thinly papyraceous laminae; pinnules with toothed margins ending in an acute apex; sori in 1 or 2 rows on each side of pinnule costa; indusia orbicular-reniform, membranaceous, margin eroded (Zhang et al. 2013). The main morphological characters of these four species of Dryopteris are presented in the table below.

Discussion

The data obtained in this study allowed us to suggest a more accurate view of the taxonomic position of D. blanfordii subsp. nigrosquamosa. Our results demonstrated that D. laeta and D. blanfordii subsp. nigrosquamosa belong to the same clade as D. arguta and D. marginalis. According to the classification system of the genus Dryopteris by Fraser-Jenkins (1986), D. arguta and D. marginalis belong to sect. Pallidae, while D. blanfordii belongs to sect. Remotae. More recent classifications divide the genus Dryopteris into either five (Sessa et al. 2012b) or 13 different clades (Zhang et al. 2012). We have concluded that D. blanfordii subsp. nigrosquamosa together with D. laeta, D. arguta and D. marginalis belong to the Dryopteris clade following Zhang et al. (2012) or clade I according to Sessa et al. (2012b). Dryopteris arguta and D. marginalis are closely related American species (from western and eastern parts of North America respectively) and the D. laeta specimen is from a population located in Iwaizumi (Iwate prefecture, Japan). According to some recent studies (Widén et al. 2015), Dryopteris goeringiana (Kunze) Koidz. is a synonym of D. laeta, growing in Japan. Our analyses rather showed that D. goeringiana proves to be related to Dryopteris stewartii Fraser-Jenk., Dryopteris lacera (Thunb.) Kunze and Dryopteris sieboldii (Van Houtte ex Mett.) Kuntze (Fig. 2), which agrees with the data of other authors (Sessa et al. 2012b). At the same time, D. laeta from Japan does not belong to this group of Dryopteris species (Ebihara 2011, Zhang et al. 2012) (Fig. 2). The relationship between species from Europe, Central America and Asia may indicate that a long-distance dispersal event occurred. Unfortunately, it is impossible to indicate where the ancestor of these species might have originated.

Figure 2. 

Topology from ML and BI analyses of Dryopteris species using six marker regions consisting of 3734 total bases (psbA-trnH, rbcL-accD, rbcL, trnL-trnF, trnP-petG, rps4-trnS and matK). The tree is drawn to scale, with branch lengths measured in the number of substitutions per site. Bootstrap values are specified at the branch nodes (cut off >50%) / Bayesian РР. Dryopteris blanfordii subsp. nigrosquamosa and closely related species are marked with a blue rectangle.

Dryopteris arguta, D. marginalis, D. laeta and D. blanfordii subsp. nigrosquamosa differ from each other in many morphological characters (Table 1), but share the structure of segments of second and third orders. The second order segments (pinnules) are short- or very short-stalked (1–5(10) mm). The distal part of the pinnule is attenuated and elongated, with its distal third alate and lacking sori. The basal basiscopic to acroscopic pinnulae length ratio is 1–1.5. The third order segments, when present, are isomorphic. Pinnules are serrate, ending with more or less prominent spiny teeth.

Conclusion

The relationship of D. blanfordii subsp. nigrosquamosa within the Dryopteris genus was defined using phylogenetic analyses based on chloroplast markers. Our results demonstrate that Asian species D. blanfordii subsp. nigrosquamosa and D. laeta belong to the same clade as two North American species D. arguta and D. marginalis, with all four species being part of the Dryopteris section Dryopteris.

Acknowledgements

This investigation (except sequencing works) was supported by the Russian Foundation for Basic Research, grant no. 14-04-01852а. The sequencing works were supported by Russian Science Foundation grant no. 14-50-00029.

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Appendix 1

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GenBank accession numbers of marker sequences of Dryopteris species included in the study. All sequences of D. blanfordii subsp. nigrosquamosa were newly generated in this study.

Species matK GenBank accession/ voucher or isolate psbA-trnH GenBank accession / voucher or isolate rbcL-accD GenBank accession / voucher or isolate rps4-trnS GenBank accession / voucher or isolate trnL-trnF GenBank accession / voucher or isolate trnP-petG GenBank accession / voucher or isolate rbcL GenBank accession / voucher or isolate
Dryopteris abbreviata Kuntze JN189448.1 / isolate E335 JN189664.1 / isolate E335 JN189231.1 / isolate E335 JN189126.1 / - JN189342.1 / isolate E335 JN189557.1 / isolate E335
Dryopteris aemula (Ait.) Kuntze JN189407.1 / isolate E113 JN189625.1 / isolate E113 JN189189.1 / isolate E113 AY268816.1 / - JN189301.1 / isolate E113 AY268881.1 / -
Dryopteris affinis (Lowe) Fraser-Jenk. JN189408.1 / isolate E114 JN189626.1 / isolate E114 JN189190.1 / isolate E114 AY268780.1 / - JN189302.1 / isolate E114 AY268849.1 / -
Dryopteris alpestris Tagawa ex Ching & S.K.Wu JQ941627.1/ Heng 32147 (UC) JN189428.1 / isolate E314 JN189645.1 / isolate E314 JN189210.1 / isolate E314 JX535868.1 / Xiaohua Jin & Liang Zhang 11103 (CDBI) JN189322.1 / isolate E314 JX535858.1 / Xiaohua Jin & Liang Zhang 11103 (CDBI)
Dryopteris antarctica (Baker) C.Chr. JQ941648.1/ Hennequin 2009-R109 (REU) JN189467.1 / isolate E378 JN189682.1 / isolate E378 JN189250.1 / isolate E378 JN189141.1 / isolate E378 JN189356.1 / isolate E378 JN189577.1 / isolate E378
Dryopteris aquilinoides (Desv.) C.Chr. JQ941617.1/ Kessler 13855 (UC) JN189429.1 / isolate E315 JN189646.1 / isolate E315 JN189211.1 / isolate E315 JN189106.1 / isolate E315 AY268803.1/ Ranker 1536 JN189323.1 / isolate E315 JN189537.1 / isolate E315 AY268868.1/ -
Dryopteris arguta (Kaulf.) Maxon JQ941647.1/ EBS 35 (WIS) JQ941660.1/ EBS 36 (WIS) JN189400.1 / isolate E069 JN189619.1 / isolate E069 JQ936838.1 / EBS 36 (WIS) AY278397.1 / - JN189294.1 / isolate E069 JN189509.1 / isolate E069 JQ935258.1/ EBS 36 (WIS)
Dryopteris ardechensis Fraser-Jenk. JN189487.1 / isolate NH56 JN189702.1 / isolate NH56 JN189271.1 / isolate NH56 AY268817.1 / - JN189377.1 / isolate NH56 JN189596.1 / isolate NH56
Dryopteris assimilis S.Walker JQ941622.1/ Skvortsov 1.VIII.1982 (NY) JN189409.1 / isolate E117 JN189627.1 / isolate E117 JN189191.1 / isolate E117 JN189086.1 / isolate E117 JN189303.1 / isolate E117 JN189517.1 / isolate E117
Dryopteris austriaca (Jacq.) Woyn. ex Schinz & Thell. JQ941637.1/ Degn 25 (NY) JN189410.1 / isolate E119 JN189628.1 / isolate E119 JN189192.1 / isolate E119 JN189087.1 / isolate E119 JN189304.1 / isolate E119 JN189518.1 / isolate E119
Dryopteris barbigera (Moore) Kuntze JN189431.1 / isolate E317 JN189647.1 / isolate E317 JN189213.1 / UC<USA-CA> Miehe 94-191-14 JN189108.1 / isolate E317 JN189325.1 / isolate E317 JN189539.1 / UC<USA-CA> Miehe 94-191-14
Dryopteris bissetiana (Baker) C.Chr. AB575740.1 / TNS:763335 JN189693.1 / isolate NH28 DQ191829.1 / SG Lu/C75 JN189261.1/ isolate NH28 AY268796.1 / R. Moran, COLO JN189367.1 / isolate NH28 AY268862.1 / -
Dryopteris campyloptera (Kunze) Clarkson JQ941638.1/ EBS 22 (WIS) JQ941619.1/ EBS 19 (WIS) JN189395.1 / isolate E058 JN189614.1 / isolate E058 JQ936819.1 / EBS 19 (WIS) AY268801.1 / WJ Cody 23484 FR731970.1/ isolate A183 JN189289.1 / isolate E058 JQ935255.1 / EBS 19 (WIS) AY268866.1/ -
Dryopteris carthusiana (Vill.) H.P.Fuchs JQ941634.1/ EBS 42 (WIS) JQ941655.1/ EBS 43 (WIS) JQ941653.1/ EBS 7 (WIS) JQ941640.1/ EBS 41 (WIS) JN189402.1 / isolate E075 JN189621.1 / isolate E075 JN189184.1 / isolate E075 JQ936839.1/ EBS 42 (WIS) AY268777.1 / GW Argus 9327, COLO JN189296.1 / isolate E075 JQ683076.1/ EBS 43 (WIS) JQ683064.1/ EBS 7 (WIS) JN189511.1 / isolate E075 JQ935266.1/ EBS 7 (WIS) JQ935272.1/ EBS 43 (WIS)
Dryopteris caucasica (A.Braun) Fraser-Jenk. & Corley JQ941604.1/ Christenhusz 4309 (UC) JN189432.1 / isolate E318 JN189648.1 / isolate E318 JN189214.1 / isolate E318 JN189109.1 / isolate E318 JN189326.1 / isolate E318 JN189540.1 / isolate E318
Dryopteris celsa (W.Palmer) Small JQ941652.1/ EBS 27 (WIS) JQ941658.1/ Price 94-2 (NY) JQ936652.1 / Price 94-2 (NY) JN189609.1 / isolate E043 JN189175.1 / isolate E043 JQ936822.1/ EBS 49 (WIS) JN105314.1 / EBS49 (WIS) JQ683075.1 / Price 94-2 (NY) JQ935249.1 / EBS 49 (WIS)
Dryopteris championii (Benth.) C.Chr. JN189480.1 / isolate NH31 AB575742.1/ TNS:764357 JN189694.1 / isolate NH31 DQ151856.1 / SG Lu/M3 AY268797.1 / R. Moran, COLO KC896581.1/ isolate CHA 1419 JN189368.1 / isolate NH31 AY268863.1 / - KC896547.1/ isolate CHA 1419
Dryopteris chinensis Koidz. AB575743.1 / TNS:763933 JN189649.1 / isolate E319 JN189215.1 / isolate E319 JX535819.1/ Liang Zhang & Zhangming Zhu 1114 (CDBI) JX535872.1 / Liang Zhang & Zhangming Zhu 1114 (CDBI) JN189327.1 / isolate E319 AB575119.1 / TNS:763933
Dryopteris chrysocoma (Christ) C.Chr. JN189434.1 / isolate E320 JN189650.1 / isolate E320 JN189216.1 / isolate E320 DQ191832.1/ SG Lu/C76 DQ514495.1 / LJM079 JN189328.1 / isolate E320 DQ508773.1 / -
Dryopteris cinnamomea C.Chr. JN189420.1 / isolate E279 JN189638.1 / isolate E279 JN189202.1 / isolate E279 AY278398.1 / - FR731991.1/ A190 JN189314.1 / isolate E279 JN189528.1 / isolate E279
Dryopteris clintoniana (D.C.Eaton) Dowell JQ941605.1/ EBS 16 (WIS) JQ941626.1/ EBS 8 (WIS) JQ936651.1 / EBS 8 (WIS) JN189389.1/ isolate E020 JN189608.1 / isolate E020 JN189174.1 / isolate E020 JQ936813.1/ EBS 8 (WIS) JQ683004.1 / EBS 8 (WIS) JN189283.1 / isolate E020 JQ935247.1 / EBS 8 (WIS) KF186502.1/ OAC 96815
Dryopteris crispifolia Rasbach, Reichst. & G.Vida JQ941650.1 / BPSSE JN189488.1 / isolate NH58 JN189703.1 / isolate NH58 JN189272.1 / isolate NH58 AY268819.1 / - JN189378.1 / isolate NH58 AY268884.1 / -
Dryopteris costalisora Tagawa JN189493.1 / isolate NH85 JN189710.1 / isolate NH85 JN189278.1 / isolate NH85 JN189170.1 / isolate NH85 JN189384.1 / isolate NH85 JN189603.1 / isolate NH85
Dryopteris crassirhizoma Nakai AB575746.1 / TNS:764333 JN189651.1 / isolate E321 JN189217.1 / isolate E321 AY268805.1 / - AY278399.1/ - JN189329.1 / isolate E321 JN189543.1 / isolate E321 AY268870.1/ - KC896537.1/ isolate CRA 176
Dryopteris cristata (L.) A.Gray JQ941613.1/ EBS 58 (WIS) JQ941608.1/ EBS 68 (WIS) JQ941614.1/ Montgomery 07-99 (NY) JQ941609.1/ Leoschke 2119 (NY) JQ941621.1/ ESB 26 (WIS) JQ936659.1 / EBS 68 (WIS) JQ936654.1/ Leoschke 2119 (NY) JQ947928.1 / EBS 58 (WIS) JQ936835.1 / Montgomery 07-99 (NY) JQ682997.1 / EBS 68 (WIS) JN189330.1 / isolate E322 JQ683056.1/ Montgomery 07-99 (NY) JN189299.1/ isolate E089 JQ683055.1/ ESB 26 (WIS) JQ683065.1/ EBS 68 (WIS) JQ683070.1/ EBS 58 (WIS) JQ683073.1/ EBS 52 (WIS) JQ683078.1/ Leoschke 2119 (NY) JQ935251.1 / Montgomery 07-99 (NY) KF186503.1/ OAC 96818
Dryopteris cycadina (Franch. & Sav.) C.Chr. JN189436.1 / isolate E322 JN189652.1 / isolate E322 JN189218.1 / isolate E322 DQ191835.1/ SG Lu/C38 JN189113.1 / isolate E322 AY278400.1/ - JN189330.1 / UC<USA-CA> RBCTW-078 JN189544.1 / isolate E322 EF463127.1/ - AY587115.1/ -
Dryopteris cystolepidota (Miq.) C.Chr. JN189485.1 / isolate NH52 JN189699.1 / isolate NH52 JN189268.1 / isolate NH52 AY268813.1 / - JN189374.1 / isolate NH52 JN189593.1 / isolate NH52 AY268878.1/ -
Dryopteris dilatata (Hoffm.) A.Gray JQ941646.1/ Hennequin 2010-B1 (P) JQ941599.1/ Schuettpelz 535 (DUKE) JQ941645.1/ Camoletto 2021 (NY) JN189465.1 / isolate E375 JQ936648.1/ Schuettpelz 535 (DUKE) JQ936650.1/ Camoletto 2021 (NY) JN189680.1 / isolate E375 JQ936841.1 / Schuettpelz 535 (DUKE) JQ683001.1 / Schuettpelz 535 (DUKE) AY268779.1/ Kasnoborov 679, COLO JN189354.1 / isolate E375 JN189575.1 / - AY268848.1/ - JQ935276.1/ Schuettpelz 535 (DUKE)
Dryopteris dickinsii (Franch. & Sav.) C.Chr. JN189489.1 / isolate NH59 JN189704.1 / isolate NH59 JN189273.1 / isolate NH59 DQ191839.1/ SG Lu/QC3 JX535875.1 / K. Ohora s.n.; VS766491 (TNS) AY268820.1/ - JN189379.1 / isolate NH59 JN189598.1 / isolate NH59 AB575125.1/ TNS:766491 DDU05622.1/ -
Dryopteris dehuaensis Ching & K. H. Shing DQ191838.1/ SG Lu/O9
Dryopteris erythrosora (D. C. Eat.) O. Kuntze KM208819.1 / - PSMT3776/ - EF590692.1/ NMNH 06-8357 JN189687.1 / isolate NH13 JN189255.1 / isolate NH13 DQ191840.1/ SG Lu/B33 AY268787.1 / Geiger 94, COLO JN189361.1 / isolate NH13 AY268852.1 / - AB232392.1/ Tsutsumi CT1001 AY587111.1/ - KC896551.1/ isolate ERY 2006899
Dryopteris expansa (C. Presl) Fraser-Jenk. & Jermy JQ941606.1/ EBS 37 (WIS) JQ941612.1/ EBS 40 (WIS) JQ941633.1/ EBS 30 (WIS) JQ941610.1/ EBS 33 (WIS) AB575750.1 / TNS:765180 JN189616.1 / isolate E064 JN189180.1 / isolate E064 KF020383.1/ Nelson 7921 AY268775.1 / Nelson 7921 JN189291.1 / isolate E064 JQ935275.1 / EBS 37 (WIS) EF463179.1/ Christenhusz 4263 (TUR) AB575127.1/ TNS:765180 AB908270.1/ -
Dryopteris filix-mas (L.) Schott JQ941618.1/ EBS 38 (WIS) JQ941611.1/ EBS 32 (WIS) JN189398.1 / isolate E066 JN189617.1 / isolate E066 JN189181.1 / isolate E066 FR731980.1 / isolate A213 JQ683060.1 / EBS 38 (WIS) JN189507.1 / isolate E066 JQ935265.1/ EBS 38 (WIS) KM114199.1/ Lehtonen 725 (TUR) JF832067.1/ -
Dryopteris flaccisquama A.Rojas JN189411.1 / isolate E236 JN189629.1 / isolate E236 JN189193.1 / isolate E236 JN189088.1 / isolate E236 JN189305.1 / isolate E236 JN189519.1 / -
Dryopteris fragrans (L.) Schott JQ941603.1/ EBS 47 (WIS) JN189403.1/ isolate E081 JN189185.1/ isolate E081 AY268800.1/ Kelso 83-221 JQ683049.1/ EBS 53 (WIS) JN189297.1/ isolate E081 AB575129.1/ TNS:743728 JN189512.1 / isolate E081 JQ935274.1/ EBS 53 (WIS) AY268865.1/ -
Dryopteris formosana (Christ) C.Chr. AB575751.1 / TNS:763153 JN189653.1 / isolate E323 JN189219.1 / isolate E323 AY268793.1 / R. Moran, COLO JN189331.1 / isolate E323 JN189545.1 / - AB575128.1/ TNS:763153 AY268857.1/ -
Dryopteris fuscipes C.Chr. AB575752.1/ TNS:762541 DQ191841.1/ SG Lu/M2 KC896583.1/ FUS 1058 AB575130.1/ TNS:762541 KC896549.1 \ isolate FUS 1058
Dryopteris future A.R.Sm. JN189426.1 / isolate E299 JN189643.1 / isolate E299 JN189208.1 / isolate E299 JN189103.1 / isolate E299 JN189320.1 / isolate E299 JN189534.1 / -
Dryopteris goldiana (Hook.) A.Gray JN189396.1 / isolate E063 JN189615.1 / isolate E063 JN189179.1 / - FR731984.1 / isolate A202 JN189290.1 / isolate E063 AF537228.1 / -
Dryopteris goeringiana (G. Kze.) Koidz. JN189474.1 / isolate NH16 JN189688.1 / isolate NH16 JN189256.1 / - AY268790.1 / - AF515241.1/ - JN189362.1 / isolate NH16 AY268855.1 / R. Moran; COLO», cultivated in NY botanical garden KC896541.1/ isolate GOE 080368
Dryopteris gymnosora (Mak.) C. Chr. JN189438.1 / isolate E324 JN189654.1 / isolate E324 JN189220.1 / isolate E324 JX535824.1/ - JN189115.1 / isolate E324 JX535877.1/ - JN189332.1 / isolate E324 AB575132.1 / TNS:763325
Dryopteris guanchica Gibby & Jermy JN189463.1 / isolate E373 JN189678.1 / isolate E373 JN189246.1 / - FR731992.1 / isolate A207 JN189352.1 / isolate E373 JN189573.1 / isolate E373
Dryopteris hawaiiensis (Hillebr.) Robinson JN189470.1 / isolate MA85 JN189685.1 / isolate MA85 JN189253.1 / isolate MA85 JN189144.1 / isolate MA85 AY268784.1/ Geiger 74, COLO JN189359.1 / isolate MA85 AY268840.1 / -
Dryopteris hondoensis Koidz. AB575761.1 / TNS:764343 JN189689.1 / isolate NH17 JN189257.1 / isolate NH17 AY268791.1 / R. Moran, COLO JN189363.1 / isolate NH17 JN189583.1 / isolate NH17 AB575139.1/ TNS:764343
Dryopteris inaequalis (Schlecht.) Kuntze JN189440.1 / isolate E326 JN189655.1 / isolate E326 JN189222.1 / isolate E326 JN189117.1 / isolate E326 JN189333.1 / isolate E326 JN189548.1 / isolate E326
Dryopteris intermedia (Willd.) A.Gray JQ941657.1/ EBS 63 (WIS) JQ941659.1/ EBS 44 (WIS) JQ941642.1/ EBS 69 (WIS) JQ941663.1/ EBS 13 (WIS) JQ941624.1/ ESB 15 (WIS) JQ941630.1/ EBS 18 (WIS) JQ936670.1 / EBS 63 (WIS) JN189613.1 / isolate E056 JN189178.1 / isolate E056 FR731994.1 / Alanko 96215 (H:1695737) FR731993.1/ isolate A141 JN189288.1 / isolate E056 AB575143.1 / TNS:766638
Dryopteris juxtaposita Christ JN189441.1 / isolate E327 JN189656.1 / isolate E327 JN189223.1 / isolate E327 DQ191848.1/ SG Lu/66 JN189118.1 / isolate E327 AY268810.1/ - JN189334.1 / isolate E327 JN189549.1 / isolate E327 AY268875.1/ -
Dryopteris karwinskyana (Mett.) Kuntze JN189413.1 / isolate E241 JN189631.1 / isolate E241 JN189195.1 / isolate E241 JN189090.1 / isolate E241 JN189307.1 / isolate E241 JN189521.1 / isolate E241
Dryopteris komarovii Kossinsky JN189442.1 / isolate E328 JN189657.1 / isolate E328 JN189224.1 / isolate E328 JN189119.1 / isolate E328 JN189335.1 / isolate E328 JN189550.1 / -
Dryopteris knoblochii A. R. Sm. JN189414.1 / isolate E242 JN189632.1 / isolate E242 JN189196.1 / isolate E242 JN189091.1 / isolate E242 JN189308.1 / isolate E242 JN189522.1 / -
Dryopteris kobayashii Kitag. AB908374.1/ haplotype: Type 6
Dryopteris labordei (Christ) C.Chr. JN189492.1 / isolate NH84 JN189709.1 / isolate NH84 JN189277.1 / isolate NH84 JN189169.1 / isolate NH84 JN189383.1 / isolate NH84 JN189602.1 / isolate NH84 EF463128.1/ -
Dryopteris lacera (Thunb.) Ktze. AB575770.1 / TNS:774850 JN189691.1 / isolate NH23 JN189259.1 / isolate NH23 DQ191851.1/ SG Lu/QC36 AY268794.1 / R. Moran, COLO KC896572.1/ isolate LAC 1027 JN189365.1 / isolate NH23 JN189585.1 / isolate NH23 KC896538.1/ isolate LAC 1027
Dryopteris ludoviciana (Kze.) Small JQ941656.1/ EBSlud2 (WIS) JQ941628.1/ EBSlud3 (WIS) JQ941639.1/ EBSlud4 (WIS) JQ941654.1/ EBS 48.A (WIS) JQ936667.1 / EBSlud2 (WIS) JQ947929.1 / EBSlud4 (WIS) JN189186.1 / isolate E082 JQ936837.1/ EBS 48 (WIS) JQ682979.1 / EBSlud2 (WIS) JN189298.1 / isolate E082 JQ935277.1 / EBSlud3 (WIS) JQ935273.1/ EBS 48 (WIS)
Dryopteris marginalis (L.) A.Gray JQ941607.1/ EBS 17 (WIS) JN189393.1 / isolate E055 JN189612.1 / isolate E055 JN189177.1 / isolate E055 FR731986.1 / isolate A203 JN189287.1 / isolate E055 EF463181.1 / - KF186511.1/ OAC 96835
Dryopteris maxonii Underw. & C.Chr. JN189421.1 / isolate E284 JN189639.1 / isolate E284 JN189203.1 / isolate E284 JN189098.1 / isolate E284 JN189315.1 / isolate E284 JN189529.1 / isolate E284
Dryopteris medioxima Koidz. AB575774.1/ TNS:776976 AB575152.1/ 776976
Dryopteris munchii A.R.Sm. JQ941632.1/ EBS 54 (WIS) JN189427.1 / isolate E311 JN189644.1 / isolate E311 JN189209.1 / - JQ936827.1/ Hoshizaki (UC) AY268822.1 / - JN189321.1 / WIS EBS54 JN189535.1 / isolate E311
Dryopteris odontoloma (Moore) C. Chr. JN189483.1 / isolate NH46 JN189696.1 / isolate NH46 JN189265.1 / - DQ191859.1/ - AY268807.1 / - JN189371.1 / isolate NH46 JN189590.1 / isolate NH46 AY268872.1/ -
Dryopteris oligodonta (Desv.) Pichi-Serm. JQ941643.1/ voucher Hennequin 2010-C11 (P) JN189464.1 / isolate E374 JN189679.1 / isolate E374 JN189247.1 / isolate E374 FR731987.1 / isolate A156 JN189353.1 / isolate E374 JN189574.1 / isolate E374
Dryopteris pacifica (Nakai) Tag. AB575778.1 / TNS:763312 JN189700.1 / isolate NH53 JN189269.1 / isolate NH53 DQ191860.1/ - JX535838.1/ - JX535891.1 / - KC896588.1/ isolate PAC 487 AY268814.1/ - JN189375.1 / isolate NH53 JN189594.1 / - AB575157.1/ TNS:763312
Dryopteris panda (C. B. Cl.) C. Chr. JN189443.1 / isolate E329 JN189658.1 / isolate E329 JN189225.1 / - DQ191861.1/ - JN189120.1 / isolate E329 JN189336.1 / isolate E329 JN189551.1 / -
Dryopteris patula (Sw.) Underw. JN189391.1 / isolate E047 JN189610.1 / isolate E047 KJ464668.1 / isolate MS1104 AY268823.1 / - JN189285.1 / isolate E047 KJ464427.1 / - AY268888.1 -
Dryopteris pentheri (Krasser) C. Chr. JN189466.1 / isolate E377 JN189681.1 / isolate E377 JN189249.1 / isolate E377 JN189140.1 / isolate E377 JN189355.1 / isolate E377 KF992481.1 / -
Dryopteris polita Rosenst. KJ196593.1/ S.Y. Dong 718 AB575779.1 / TNS:763901 JN189713.1 / isolate NH92 JN189281.1 / isolate NH92 KJ196700.1 / S.Y. Dong 718 JN189387.1 / isolate NH92 AB575158.1 / TNS:763901
Dryopteris polylepis (Franch. & Sav.) C. Chr. AB575780.1 / TNS:764340 JN189695.1 / isolate NH33 JN189263.1 / isolate NH33 AY268798.1 / R. Moran, COLO JN189369.1 / isolate NH33 AB575159.1 / TNS:764340
Dryopteris pseudofilix-mas (Fée) Rothm. JN189415.1 / isolate E259 JN189633.1 / isolate E259 JN189197.1 / isolate E259 AY278404.1 / - JN189309.1 / isolate E259 JN189523.1 / -
Dryopteris pulcherrima Ching JN189406.1 / isolate E108 JN189624.1 / isolate E108 JN189188.1 / isolate E108 JN189083.1 / isolate E108 AY268811.1/ - JN189300.1 / isolate E108 JN189515.1 / isolate E108
Dryopteris pycnopteroides (Christ) C. Chr. AB575781.1 / TNS:764373 JN189690.1 / isolate NH22 JN189258.1 / isolate NH22 DQ191868.1/ SG Lu/C79 AY268799.1 / R. Moran, COLO JN189364.1 / isolate NH22 AB575160.1 / TNS:764373 AY587114.1/ -
Dryopteris remota Hayata JQ941635.1/ Schuettpelz 528 (DUKE) JQ941616.1/ Moran.A (COLO) JQ936655.1 / Moran (COLO) JN189640.1 / isolate E285 JN189204.1 / Isolate E285 JQ936826.1/ Moran (COLO) JQ682983.1 / Moran (COLO) AY268792.1/ R. Moran, COLO JN189316.1 / isolate E285 JQ683062.1/ Moran (COLO) AY268858.1 / - JQ935253.1/ Moran (COLO)
Dryopteris reflexosquamata Hayata JN189494.1 / isolate NH86 JN189711.1 / isolate NH86 JN189279.1 / isolate NH86 JN189171.1 / isolate NH86 JN189385.1 / isolate NH86 JN189604.1 / isolate NH86
Dryopteris rosea (E.Fourn.) Mickel & Beitel JN189416.1 / isolate E260 JN189634.1 / isolate E260 JN189198.1 / isolate E260 JN189093.1 / isolate E260 JN189310.1 / isolate E260 JN189524.1 / -
Dryopteris rossii C.Chr. JN189423.1 / isolate E286 JN189641.1 / isolate E286 JN189205.1 / isolate E286 JN189100.1 / isolate E286 JN189317.1 / isolate E286 JN189531.1 / isolate E286
Dryopteris sacrosancta Koidz. AB575784.1 / TNS:764377 JN189698.1 / isolate NH51 JX535843.1 / cult. Tsukuba Botanical Garden; VS764377 (TNS) JN189267.1/ AFSSE JX535896.1 / cult. Tsukuba Botanical Garden; VS764377 (TNS) AY268812.1/ - JN189373.1 / isolate NH51 AY268877.1 / - AB575163.1/ TNS:764377
Dryopteris salvinii (Bak.) Kuntze JN189418.1 / isolate E264 JN189636.1 / isolate E264 JN189200.1 / isolate E264 JN189095.1 / isolate E264 JN189312.1 / isolate E264 JN189526.1 / isolate E264
Dryopteris scottii (Bedd.) Ching JQ941623.1/ voucher RBC 202 (UC) JN189444.1 / isolate E330 JN189659.1 / isolate E330 JN189226.1 / iIsolate E330 DQ191872.1/ SG Lu/B31 JX535898.1 / Hai He 4 (CTC) DQ514498.1/ MMO03-313 JN189337.1 / isolate E330 JX535863.1 / - DQ508775.1/ -
Dryopteris simplicior Mickel & Beitel JN189419.1 / isolate E265 JN189637.1 / isolate E265 JN189201.1 / isolate E265 JN189096.1 / isolate E265 JN189313.1 / isolate E265 JN189527.1 / isolate E265
Dryopteris sordidipes Tagawa JN189495.1 / isolate NH88 AB575793.1/ TNS:763050 JN189712.1 / isolate NH88 JN189280.1 / isolate NH88 JX535848.1/ S. Tagane & K. Fuse TF009; VS763050 (TNS) JX535902.1 / S. Tagane & K. Fuse TF009; VS763050 (TNS) JN189386.1 / isolate NH88 AB575172.1 / TNS:763050
Dryopteris stenolepis (Baker) C.Chr. JN189445.1 / isolate E331 JN189660.1 / isolate E331 JN189227.1 / isolate E331 DQ191877.1/ SG Lu/B28 JN189122.1 / isolate E331 AY268824.1/ - JN189338.1 / isolate E331 AY268889.1 / -
Dryopteris stewartii Fraser-Jenk. JN189478.1 / isolate NH26 JN189692.1 / isolate NH26 JN189260.1 / isolate NH26 AY278405.1 / - JN189366.1 / isolate NH26 JN189586.1 / isolate NH26
Dryopteris subbipinnata W.H.Wagner & R.W.Hobdy JN189469.1 / isolate MA116 JN189684.1 / isolate MA116 JN189252.1 / isolate MA116 AY268765.1 / Oppenheimer H50074, COLO JN189358.1 / isolate MA116 JN189579.1 / isolate MA116
Dryopteris sublacera Christ JN189446.1 / isolate E332 JN189661.1 / isolate E332 JN189228.1 / isolate E332 DQ191878.1/ SG Lu/59 JN189123.1 / isolate E332 AY268788.1/ Geiger 95, COLO DQ514501.1/ LJM080 JN189339.1 / isolate E332 JN189554.1 / isolate E332
Dryopteris tokyoensis (Matsum.) C.Chr. JQ941651.1/ voucher Moran.B (COLO) AB575795.1 / TNS:766452 JN189683.1 / isolate JGtok JN189251.1 / isolate JGtok AY268795.1 / R. Moran, COLO JN189357.1 / isolate JGtok AB575174.1 / TNS:766452
Dryopteris uniformis Makino AB575797.1 / TNS:774834 JN189662.1 / isolate E333 DQ191883.1 / - AY268806.1 / - JN189340.1 / isolate E333 JN189555.1 / isolate E333
Dryopteris varia (L.) Kuntze AB575798.1 / TNS:763911 JN189663.1 / isolate E334 JN189230.1 / - JX535852.1/ - AY736355.1 / - JN189341.1 / isolate E334 AB575178.1 / TNS:763911
Dryopteris wallichiana (Spreng.) Hyl. JN189388.1 / - JN189607.1 / isolate E001 DQ191884.1 / - AY268761.1 / - JN189282.1 / isolate E001 AY268826.1 / - KF992482.1/ S. Hennequin 288 (P REU)
Dryopteris xanthomelas (Christ) C.Chr. JN189449.1 / - JN189665.1 / isolate E337 JN189232.1 / - JN189127.1 / - JN189343.1 / isolate E337 JN189558.1 / isolate E337
Dryopteris blanfordii (C.Hope) C.Chr. subsp. nigrosquamosa (Ching) Fraser-Jenk. KT876448.1/ - KT876440.1 / - KT876441.1 / - KT876442.1 / - KT876443.1 / - KT876444.1 / - KT876447.1 / -
Polystichum andersonii Hopkins JQ941662.1/ EBS 39 (WIS) JN189401.1 / isolate E073 JN189620.1 / isolate E073 JN189183.1 / isolate E073 JN189078.1 / isolate E073 JN189295.1 / isolate E073 JN189510.1 / isolate E073
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