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Molecular Phylogeny of the Fern Family Dryopteridaceae Inferred

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Molecular Phylogeny of the Fern Family Dryopteridaceae Inferred Molecular Phylogeny of the Fern Family Dryopteridaceae Inferred from Chloroplast rbc L and atp B Genes Author(s): Hong‐Mei Liu, Xian‐Chun Zhang, Wei Wang, Yin‐Long Qiu, and Zhi‐Duan Chen Source: International Journal of Plant Sciences, Vol. 168, No. 9 (November/December 2007), pp. 1311-1323 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/521710 . Accessed: 02/04/2015 02:30 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. http://www.jstor.org This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 02:30:39 AM All use subject to JSTOR Terms and Conditions Int. J. Plant Sci. 168(9):1311–1323. 2007. Ó 2007 by The University of Chicago. All rights reserved. 1058-5893/2007/16809-0007$15.00 DOI: 10.1086/521710 MOLECULAR PHYLOGENY OF THE FERN FAMILY DRYOPTERIDACEAE INFERRED FROM CHLOROPLAST rbcL AND atpB GENES Hong-Mei Liu,*,y Xian-Chun Zhang,1,* Wei Wang,* Yin-Long Qiu,z and Zhi-Duan Chen2,* *State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan, Beijing 100093, People’s Republic of China; yGraduate University of Chinese Academy of Sciences, Beijing 100039, People’s Republic of China; and zDepartment of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan 48109-1048, U.S.A. Dryopteridaceae are a large family of leptosporangiate ferns exhibiting a rich diversity of morphological characteristics and generating many taxonomical controversies. Previous efforts to systematize this large group without a cladistic phylogenetic framework have not reached a consensus on circumscription of the family. In order to circumscribe the family and identify major groups within it, a broadscale phylogenetic analysis was conducted on these ferns plus representatives of putatively related families within the Eupolypods I clade. Two chloroplast loci, rbcL and atpB, were used in this study. A combined analysis of the two genes generated a well-resolved and strongly supported overall phylogeny of the Dryopteridaceae. The results indicate that the Dryopteridaceae form a monophyletic group with the exception of Didymochlaena, Hypodematium, and Leucostegia and are sister to a large clade comprising Lomariopsidaceae, Tectariaceae, Polypodiaceae, Davalliaceae, and Oleandraceae. Of the three previously unincorporated genera, Leptorumohra and Phanero- phlebiopsis are nested within the genus Arachniodes, and Diacalpe is closely related to Acrophorus. The genus Pleocnemia, previously classified in Tectariaceae, clearly belongs in Dryopteridaceae and shows a close affinity to Lastreopsis. This study identifies four major lineages within the family, each strongly supported as a mono- phyletic group. Two large genera, Dryopteris and Polystichum, as traditionally defined, are shown to be polyphyletic and need to be revised in the future. Keywords: atpB, chloroplast loci, Dryopteridaceae, ferns, molecular phylogeny, rbcL. Introduction these studies were based mainly on morphological character- istics, some of which were used as diagnostic characters, Dryopteridaceae are a large and diverse family of lepto- such as indusial characters (indusiate condition vs. exindusi- sporangiate ferns that have a pantropical distribution and ate condition), dimorphism of the fertile and sterile leaves, also many temperate representatives. Members of the family and growth habit, which have been shown to be homoplastic are usually terrestrial, medium-sized plants. The plants usu- (Smith 1995). ally have creeping rhizomes, ascending or erect, with non- Recent advances in molecular systematics of ferns make it clathrate scales at apices; petioles with numerous round, possible to address long-standing questions about the classi- vascular bundles arranged in a ring; veins pinnate or forking, fication of different fern groups. In the first insight of the in- free to variously anastomosing, with or without included vein- terfamilial relationships of ferns based on rbcL sequences, lets; spores reniform, monolete, perine winged; and the basic Hasebe et al. (1995) showed that Dryopteridaceae sensu chromosome number of x¼41, rarely 40. Kramer et al. (1990) were not monophyletic, which was sub- The circumscription of Dryopteridaceae has historically sequently confirmed by a combined molecular and morpho- been controversial (Pichi Sermolli 1977; Ching 1978; Tryon logical analysis (Pryer et al. 1995). and Tryon 1982; Kramer et al. 1990). Ching (1978) treated Recently, Li and Lu (2006a) performed a molecular analy- Dryopteridaceae in a strict sense with 14 genera, 13 of which sis of 12 recognized genera and other putatively related taxa are distributed in China, while other authors defined it in to reconstruct the phylogeny of Dryopteridaceae sensu Ching a broader sense with 32–52 genera (Pichi Sermolli 1977; (1978). Their study indicated that Dryopteridaceae sensu Tryon and Tryon 1982; Kramer et al. 1990). The major con- Ching (1978), except for Cyclopeltis J. Smith, were mono- troversy among different classifications centers on the issue phyletic but should be expanded to include Acrophorus C. of whether athyrioid, tectarioid, and onocleoid as well as lo- Presl, Diacalpe Blume, Peranema D. Don, and Ctenitis C. mariopsidoid ferns should be included in the family (table 1). Chr (including Dryopsis Holttum and Edwards). Two mono- However, no previous efforts to systematize this large group phyletic groups were also identified in their study, which cor- were conducted in a phylogenetic framework. Furthermore, responded largely to two tribes recognized by Ching (1978). Although it was very useful and the earliest scheme of molec- 1 Author for correspondence; e-mail [email protected]. ular phylogeny for this group, Li and Lu’s study (2006a) was 2 Author for correspondence; e-mail [email protected]. based on a narrowly defined Dryopteridaceae (sensu Ching Manuscript received May 2007; revised manuscript received July 2007. 1978), and only a single gene was used. Other molecular 1311 This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 02:30:39 AM All use subject to JSTOR Terms and Conditions Table 1 Genera Included in Different Classification Systems of Dryopteridaceae Ching 1978 Smith et al. 2006b Pichi Sermolli 1977 Tryon and Tryon 1982 Kramer et al. 1990 Acrorumohra Acrorumohra Acrorumohra Acrorumohraa Acrorumohraa Arachniodes Arachniodes Arachniodes Arachniodesa Arachniodes Cyrtogonellum Cyrtogonellum Cyrtogonellum Cyrtogonellum Cyrtogonelluma Cyrtomidictyum Cyrtomidictyum Cyrtomidictyum Cyrtomidictyum Cyrtomidictyuma Cyrtomium Cyrtomium Cyrtomium Cyrtomium Cyrtomiuma Dryopteris Dryopteris Dryopteris Dryopteris Dryopteris Leptorumohra ? Leptorumohra Leptorumohraa Leptorumohraa Lithostegia Lithostegia Lithostegia Lithostegia Lithostegia Nothoperanema Nothoperanemaa Nothoperanema Nothoperanema Nothoperanema Phanerophlebia Phanerophlebia Phanerophlebia Phanerophlebiaa Phanerophlebiaa Phanerophlebiopsis ? Phanerophlebiopsis Phanerophlebiopsisa Phanerophlebiopsisa Polystichum Polystichum Polystichum Polystichum Polystichum Sorolepidium ? Sorolepidiuma Sorolepidiuma Sorolepidiuma Cyclopeltis Cyclopeltis Cyclopeltis Cyclopeltis Acrophorus Acrophorus Acrophorus Acrophorus Ctenitis Ctenitis Ctenitis Ctenitis Didymochlaena Didymochlaena Didymochlaena Didymochlaena Dryopolystichum Dryopolystichum Dryopolystichum Dryopolystichum Lastreopsis Lastreopsis Lastreopsis Lastreopsis Maxonia Maxonia Maxonia Maxonia Olfersia Olfersia Olfersiaa Olfersia Peranema Peranema Peranema Peranema Plecosorusa Plecosorus Plecosorusa Plecosorusa Polybotrya Polybotrya Polybotrya Polybotrya Polystichopsis Polystichopsis Polystichopsisa Polystichopsisa Rumohra Rumohra Rumohra Rumohra Stenolepia Stenolepia Stenolepia Stenolepia Stigmatopteris Stigmatopteris Stigmatopteris Stigmatopteris Cyclodium Cyclodium Cyclodium Hypodematium Hypodematium Hypodematium Papuapterisa Papuapterisa Papuapterisa Adenoderris Adenoderris Bolbitis Bolbitis Egenolfiaa Egenolfiaa Elaphoglossum Elaphoglossum Lomagramma Lomagramma Teratophyllum Teratophyllum Coveniella Coveniella Dryopsis Dryopsis Megalastrum Megalastruma Ataxipteris Leucostegia Acropelta Acropeltaa Acropeltaa Amphiblastra Amphiblastraa Amphiblastraa Atalopteris Atalopteris Atalopterisa Camptodium Camptodiuma Camptodiuma Cionidium Cionidiuma Cionidiuma Ctenitopsis Ctenitopsisa Ctenitopsisa ? Diacalpe Diacalpea Diacalpea Dictyoxiphium Dictyoxiphiuma Dictyoxiphiuma Fadyenia Fadyeniaa Fadyeniaa Hemigramma Hemigramma Hemigrammaa Heterogonium Heterogonium Heterogonium Hypoderris Hypoderris Hypoderris Luerssenia Luerssenia Luersseniaa Pleocnemia Pleocnemia Pleocnemia Pleuroderris Pleuroderrisa Pleuroderrisa Pseudotectaria Pseudotectariaa Pseudotectariaa Psomiocarpa Psomiocarpa Psomiocarpa Pteridrys Pteridrys Pteridrys Quercifilix Quercifilixa Quercifilixa This content downloaded from 159.226.100.224 on Thu, 2 Apr 2015 02:30:39 AM All use subject to JSTOR Terms and Conditions
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