TL;DR: The paraphyly of Dryopteris is confirmed and the first strong molecular evidence on the monophyly of Acrophorus, Diacalpe, Dryopsis, Nothoperanema, and Peranema is provided, which strongly suggests that the current taxonomy of this genus is in need of revision.
Abstract: The fern genus Dryopteris (Dryopteridaceae) is among the most common and species rich fern genera in temperate forests in the northern hemisphere containing 225–300 species worldwide. The circumscription of Dryopteris has been controversial and various related genera have, over the time, been included in and excluded from Dryopteris. The infrageneric phylogeny has largely remained unclear, and the placement of the majority of the supraspecific taxa of Dryopteris has never been tested using molecular data. In this study, DNA sequences of four plastid loci (rbcL gene, rps4-trnS spacer, trnL intron, trnL-F spacer) were used to reconstruct the phylogeny of Dryopteris. A total of 122 accessions are sampled in our analysis and they represent 100 species of the expanded Dryopteris including Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema. All four subgenera and 19 sections currently recognized in Dryopteris s.s. are included. One species each of Arachniodes, Leptorumohra, and Lithostegia of Dryopteridaceae are used as outgroups. Our study confirms the paraphyly of Dryopteris and provides the first strong molecular evidence on the monophyly of Acrophorus, Diacalpe, Dryopsis, Nothoperanema, and Peranema. However, all these monophyletic groups together with the paraphyletic Acrorumohra are suggested to be merged into Dryopteris based on both molecular and morphological evidence. Our analysis identified 13 well-supported monophyletic groups. Each of the 13 clades is additionally supported by morphological synapomophies and is inferred to represent a major evolutionary lineage in Dryopteris. In contrast, monophyly of the four subgenera and 15 out of 19 sections currently recognized in Dryopteris s.s is not supported by plastid data. The genera, Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema, should all be merged into Dryopteris. Most species of these genera share a short rhizome and catadromic arrangement of frond segments, unlike the sister genus of Dryopteris s.l., Arachniodes, which has anadromic arrangement of frond segments. The non-monophyly of the 19 out of the 21 supraspecific taxa (sections, subgenera) in Dryopteris strongly suggests that the current taxonomy of this genus is in need of revision. The disagreement between the previous taxonomy and molecular results in Dryopteris may be due partly to interspecific hybridization and polyplodization. More morphological studies and molecular data, especially from the nuclear genome, are needed to thoroughly elucidate the evolutionary history of Dryopteris. The 13 well-supported clades identified based on our data represent 13 major evolutionary lineages in Dryopteris that are also supported by morphological synapomophies.
TL;DR: Four unusual terpenylated acylphloroglucinols were isolated from the diethyl ether extract of the scales and rhizomes of the fern Dryopteris wallichiana and displayed moderate in vitro nematocidal activity against Nippostrongylus brasiliensis.
TL;DR: This study clearly demonstrated that the currently defined Dryopteris is paraphyletic in relation to a clade containing the monophyletic Acrophorus, Diacalpe, Nothoperanema, and Peranema.
Abstract: Dryopteris (Dryopteridaceae) is one of the largest fern genera containing about 300 or more species. Sixty-five accessions representing all four subgenera and all 19 sections of Dryopteris recognized in current classifications, more than two-thirds of the species of Acrophorus, Diacalpe, Nothoperanema, and Peranema, three species of Arachniodes, three species of Cyrtomium, one species of Phanerophlebia, and two species of Polystichum, were sampled as ingroup in this study. Two species of Polystichopsis were used as outgroups. DNA sequences of five plastid loci (psbA-trnH spacer, rbcL gene, rps4-trnS spacer, trnL intron, trnL-F spacer) were used to infer the phylogeny of Dryopteris and related genera. Our study clearly demonstrated that the currently defined Dryopteris is paraphyletic in relation to a clade containing the monophyletic Acrophorus, Diacalpe, Nothoperanema, and Peranema. Peranema is resolved as sister to the remaining three genera, followed by Nothoperanema which in turn is sister to the Acrophorus+Diacalpe clade, but this relationship received low support. Three out of the four subgenera of Dryopteris recognized in current classifications are not recovered as monophyletic. Dryopteris subg. Nothoperanema, well supported as monophyletic by molecular and morphological data, is resurrected and re-circumscribed to contain species of the currently accepted genera Acrophorus, Diacalpe, Nothoperanema, and Peranema. Twenty-seven species are recognized and classified into four sections based on molecular and morphological evidence. Included in this treatment are 25 new combinations, names, species, and statuses, and neotypification or lectotypification of 14 names. The present study further located and cited types of an additional 28 related names, many of which have been unknown to science. Keys to sections and species and distributional information are given and nomenclatural issues are discussed. This paper also synonymized and excluded 33 nomenclaturally confusing names of Dryopteris that were formerly assigned to Acrophorus or Diacalpe.
TL;DR: The epiphytic habit and distinctive long-creeping rhizome of Revwattsia appear to be autapomorphies and do not warrant its generic status, and the molecular analysis was able to reject Watts’s 1914 hypothesis of a close relationship to Polystichum.
Abstract: Revwattsia fragilis (Watts) D.L. Jones (Dryopteridaceae), originally described as a Polystichum Roth by the pioneer Australian botanist Reverend W.W. Watts in 1914, is a rare epiphytic fern endemic to northeastern Queensland, Australia. Known from only a few populations, it is restricted to tropical rainforests in the Atherton Tablelands. We used the cpDNA markers psbA-trnH, rbcL, rbcL-accD, rps4-trnS, trnG-trnR, trnL-trnF, and trnP-petG to infer the relationships of Revwattsia fragilis within Dryopteridaceae. Based on our molecular analysis, we were able to reject Watts’s 1914 hypothesis of a close relationship to Polystichum. Its closest allies are a suite of Asian Dryopteris Adans. species including D. labordei, D. gymnosora, D. erythrosora and D. cystolepidota; maintaining Revwattsia renders Dryopteris paraphyletic. The epiphytic habit and distinctive long-creeping rhizome of Revwattsia appear to be autapomorphies and do not warrant its generic status. In the course of our investigation we confirmed that polyphyly of Dryopteris is also sustained by the inclusion of Acrorumohra (H.Ito) H.Ito, Acrophorus C.Presl, Arachniodes Blume, Diacalpe Blume, Dryopsis Holttum & P.J.Edwards, and Peranema D.Don. The epithet fragilis is occupied in Dryopteris, therefore we provide the name Dryopteris wattsii nom. nov. to accommodate R. fragilis in Dryopteris.
TL;DR: Using ‘overlooked’ cultivated stocks, nuclear PgiC intron sequences are analyzed and the hypothesis of hybrid origin of the species D. shibipedis can be explained by a combination of alleles of D. kinkiensis and those ofD.
Abstract: Dryopteris shibipedis was once treated as an extinct species in the Red List (2007) by the Japanese Ministry of the Environment, but ‘rediscovered’ in the Tsukuba Botanical Garden. To clarify its origin, using ‘overlooked’ cultivated stocks we analyzed nuclear PgiC intron sequences. As the PgiC genotype of D. shibipedis can be explained by a combination of alleles of D. kinkiensis and those of D. pacifica, the hypothesis of hybrid origin of the species is supported.
TL;DR: The morphology of the Dryopteris species occurring in sub-Saharan Africa is discussed, followed by a revision of the genus in this region, and the Cape Verde Islands in the Gulf of Guinea.
Abstract: The morphology of the Dryopteris species occurring in sub-Saharan Africa is discussed. This is followed by a revision of the genus in this region, and the Cape Verde Islands in the Gulf of Guinea. Twenty-six species are recognised for the region.
TL;DR: The fern genus Dryopsis is characterized by the presence of multicellular hairs and costal and costular grooves closed near their bases and is nested in a paraphyletic Dryopteris and therefore should be viewed as part of an expanded dryopteris.
Abstract: The fern genus Dryopsis (Dryopteridaceae) is characterized by the presence of multicellular hairs and costal and costular grooves closed near their bases. Following a recent molecular analysis the monophyletic Dryopsis is nested in a paraphyletic Dryopteris and therefore should be viewed as part of an expanded Dryopteris . The current paper treats Dryopsis as a section of D. subg. Erythrovariae because both of these two taxa have bullate scales and importantly Dryopsis has been resolved as sister to D. subg. Erythrovariae sect. Erythrovariae . Twenty-two species are here recognized in D. sect. Dryopsis . Included in this treatment are nine new combinations, names and statuses, and five names are lectotypified. The present study further located and cited types of an additional 37 related names, many of which have been poorly to science. A key and distributional information of every species recognized is given and related nomenclatural issues are discussed. The nine new names, combinations and statuses include Dryopteris sect. Dryopsis , D. dulongensis , D. leiboensis , D. nidus , D. nushanensis , D. submariformis , D. wantsingshanica , D. crassirachis and D. × fauriei . The following five names are lectotypified: Ctenitis silaensis , C. tibetica , Dryopteris aureovestita , D. mariformis , and Polystichum transmorrisonensis .
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TL;DR: A new fern species, Dryopteris jiucaipingensis P. S. Luo & Li Bing Zhang is described and illustrated from northwestern Guizhou, China and is most similar to D. lunanensis (Christ) C. Chr.
Abstract: A new fern species, Dryopteris jiucaipingensis P. S. Wang, Q. Luo & Li Bing Zhang (Dryopteris Adans., sect. Hirtipedes Fraser-Jenk.; Dryopteridaceae), is described and illustrated from northwestern Guizhou, China. Dryopteris jiucaipingensis is most similar to D. lunanensis (Christ) C. Chr., but the former taxon has light brown rachis scales and occurs in alpine regions (at an elevation of 2580 m), while the latter has nearly black rachis scales and is found in montane and lowland areas (below 900 m).
TL;DR: Dryopteris liboensis from southern Guizhou, China, is described and illustrated and is unique among Chinese taxa of Dryopteris sect.
Abstract: Dryopteris liboensis P. S. Wang, X. Y. Wang & Li Bing Zhang, a new species of Dryopteris Adans. (Dryopteridaceae) from southern Guizhou, China, is described and illustrated. Dryopteris liboensis is assigned to Dryopteris subg. Erythrovariae (H. Ito) Fraser-Jenk. on the basis of its stiff and narrow petiole and rachis scales, but the new species is noteworthy within the section because these scales are not bullate. Dryopteris liboensis has stiff, narrowly lanceolate and filiform scales on the petiole and rachis and is therefore considered a member of the D. erythrosora (D. C. Eaton) Kuntze group of Dryopteris sect. Erythrosoriae H. Ito. It is unique among Chinese taxa of Dryopteris sect. Erythrovariae in having leaves 1- or 2-pinnate basally, with zero to three free lobes, and 1-pinnatifid at the blade apex.
TL;DR: The climatic and geographic conditions in Dashiwei Tiankeng Group were very suitable for the growth of fern plants,the fern resources were very rich and were important plant resources in west Guangxi Province, and the fern plant species in this area had high content of active components.
Abstract: [Objective]To investigate the medicinal fern resources of Dryopteridaceae in Dashiwei Tiankeng Group. [Method]Through field investigation,sample collection,data reference and sample identification,the medicinal fern resources of Dryopteridaceae in Dashiwei Tiankeng group were investigated,and their geographical distribution and medicinal value were introduced. [Result] Four genera of medicinal fern resources of Dryopteridaceae,which were Arachniodes Blume,Cyrtomium Presl,Dryopteris Adanson and Polystichum Roth,were found out in Dashiwei Tiankeng Group. Among the four genera,there were a total of 15 species: one species of Arachniodes Blume,which was Arachiodes assamica ( Kuhn) Ohwi; five species of Cyrtomium Presl,which were C. fortunei J. Sm,C. hookerianum ( Presl) C. Chr,C. caryotideum ( Wall. ex Hook. et Grev. ) Presl,C. balansae ( Chrit) C. Chr. and C. forunei J. Sm; five species of Dryopteris Adanson,which were Dryopteris fuscipes C. Chr, D. sieboldii ( van Hout. tEx M ett. ) O. Ktze. ,D. ryopteris varia ( L. ) O. Ktze,D. dickinsii ( Baker) C. Chr. and D. labordei ( Christ) Ching; four species of Polystichum Roth,which were Polystichum makinoi ( Tagawa) Tagawa,P. neolobatum Nakai,P. piceo-paleaceum Tagawa and P. m tsussimense ( Hook. ) J. Sm. According to their efficacy,those fern plants could be divided into several types,including the type of analgesia and clearing heat,the type of eliminating phlegm and relieving asthma,the type of detoxification and diuresis,the type of hemostasis,the type of dispelling parasites,the type of curing bone injury,the type of antivirus and antitumor,and the type of relieving convulsion,calming the liver,lowering blood pressure and relieving uneasiness. [Conclusion] The climatic and geographic conditions in Dashiwei Tiankeng Group were very suitable for the growth of fern plants,the fern resources were very rich and were important plant resources in west Guangxi Province,and the fern plant species in this area had high content of active components. Therefore,it was of great significance to protecting their biodiversity and realizing their sustainable utilization. The study provided references for the further research and analysis of medicinal and development values of Dryopteridaceae.
TL;DR: During the course of the execution of the project entitled Distribution Research of Rare Plants, the habitats of two rare ferns with obscure and/or extremely restricted distribution ranges are found, as previously unreported habitats.
Abstract: During the course of the execution of the project entitled Distribution Research of Rare Plants, we found the habitats of two rare ferns with obscure and/or extremely restricted distribution ranges, as previously unreported habitats. A subantarctic fern reported on Hamgyeong-do and Gangwon-do, Dryopteris coreano-montana Nakai, was found on Mt. Ibam in Jangseong-gun, Jeollanam-do. A subtropical fern reported south of Jeju-do, Dryopteris decipiens (Hook.) Kuntze, was newly found on Mt. Cheongwan of Jangheung-gun in Jeollanam-do.
TL;DR: The data support a classic hypothesis for reticulate evolution via allopolyploid speciation in the North America taxa, including an extinct diploid progenitor in this group, and reject deep coalescent processes such as incomplete lineage sorting in favor of more recent intercontinental hybridization and chloroplast capture.
TL;DR: New World Dryopteris are not monophyletic; vicariance has dominated the history of the North American species, while long-distance dispersal prevails in the Central and South Brazilian species, a pattern not previously seen in plants.
Abstract: Premise of the study: Dryopteris is a large, cosmopolitan fern genus ideal for addressing questions about diversifi cation, biogeography, hybridization, and polyploidy, which have historically been understudied in ferns. We constructed a highly resolved, well-supported phylogeny for New World Dryopteris and used it to investigate biogeographic patterns and divergence times. Methods: We analyzed relationships among 97 species of Dryopteris , including taxa from all major biogeographic regions, with analyses based on 5699 aligned nucleotides from seven plastid loci. Phylogenetic analyses used maximum parsimony, maximum likelihood, and Bayesian inference. We conducted divergence time analyses using BEAST and biogeographic analyses using maximum parsimony, maximum likelihood, Bayesian, and S-DIVA approaches. We explored the monophyly of subgenera and sections in the most recent generic classifi cation and of geographic groups of taxa using Templeton tests. Key results: The genus Dryopteris arose ca. 42 million years ago (Ma). Most of the Central and South American species form a well-supported clade which arose 32 Ma, but the remaining New World species are the result of multiple, independent dispersal and vicariance events involving Asia, Europe, and Africa over the last 15 Myr. We identifi ed six long-distance dispersal events and three vicariance events in the immediate ancestry of New World species; reconstructions for another four lineages were ambiguous. Conclusions: New World Dryopteris are not monophyletic; vicariance has dominated the history of the North American species, while long-distance dispersal prevails in the Central and South American species, a pattern not previously seen in plants.