Taxonomic Status of Paragymnopteris in Pteridaceae and Its Relationship with Pellaea

doi:10.7525/j.issn.1673-5102.2024.04.002

Abstract

Abstract:

The taxonomy of Cheilanthoid ferns is difficult due to morphological convergence， and there are only five species in Paragymnopteris， the taxonomy of this genus is still controversial with its members assigned to several different genera by different scholars. In this paper， the phylogenetic relationships between Paragymnopteris and other cheilanthoid fernsbased on the datasets of seven tandem chloroplast gene segments， the chloroplast genome， and the ribosomal DNA were investigated， respectively. The results showed that these three genera formed a well-supported monophyletic clade and constituted the pellaeoid group. The five species of Paragymnopteris formed two clades and each clustered with members of Pellaea. Based on the above results， this study suggested that the Pellaea should be redefined and incorporated with the Paragymnopteris into the same genus.

Key words: Paragymnopteris, Pellaea, Cheilanthoid, phylogeny, taxonomic revision, Pteridaceae

CLC Number:

S682.35

Yue YIN, Wenli YANG, Hongyu HU, Gangmin ZHANG. Taxonomic Status of Paragymnopteris in Pteridaceae and Its Relationship with Pellaea[J]. Bulletin of Botanical Research, 2024, 44(4): 491-501.

Figures/Tables 4

Table 1

Sample information of species sequenced in this study

物种 Species	凭证标本 Voucher specimen	采集地 Location	采集时间 Date
银粉背蕨Aleuritopteris argentea	张钢民 15144	四川西昌	20150818
黑柄粉背蕨A. ebenipes	张钢民 s. n.	贵州安顺	20021020
雪白粉背蕨A. niphobola	卫然 s. n.	北京房山	20201115
白边粉背蕨A. albomarginata	张钢民 457	印度	20010916
四川银旱蕨Argyrochosma connectens	张钢民 2014090	四川金川	20140818
Astrolepis sinuata	B. Bartholomew et al. 3265	墨西哥	19851021
云南旱蕨Cheilanthes bhutanica	孔宪需 6207	四川康定	19810830
旱蕨C. nitidula	卫然 201506285	四川天全	20021017
禾秆旱蕨C. tibetica	董仕勇 2061	西藏拉萨	20010815
西南旱蕨C. smithii	张钢民 256	四川康定	20010730
宜昌旱蕨C. patula	张钢民 228	重庆南川	20010724
滇西旱蕨C. brausei	张钢民 424	云南丽江	20010824
毛旱蕨C. trichophylla	张钢民 2014109	四川泸定	20140818
凤尾旱蕨C. christii	张钢民 2014107	四川泸定	20140818
C. quadripinnata	张宪春 9074	马达加斯加	20170919
C. viridis	张宪春 9172	马达加斯加	20170922
欧洲金毛裸蕨Paragymnopteris marantae_1	罗依可 2023080507	西藏吉隆	20230805
欧洲金毛裸蕨P. marantae_2	刘红梅 YN282	云南香格里拉	20041029
欧洲金毛裸蕨P. marantae_3	张钢民 2086	云南中甸	20010814
滇西金毛裸蕨P. delavayi_1	STET 1045	西藏察隅	20100721
滇西金毛裸蕨P. delavayi_2	张树仁 GGS10253	四川康定	20170718
滇西金毛裸蕨P. delavayi_3	张钢民 268	四川康定	20010801
中间金毛裸蕨P. delavayi var. intermedia_1	张宪春 4678	西藏林芝	20070929
中间金毛裸蕨P. delavayi var. intermedia_2	D.E. Boufford et.al. 35446	四川巴塘	20060630
中间金毛裸蕨P. delavayi var. intermedia_3	张宪春 8897	四川木里	20170819
中间金毛裸蕨P. delavayi var. intermedia_4	张宪春3943	西藏朗县	20060625
三角金毛裸蕨P. sargentii_1	张钢民 341	云南德钦	20010818
三角金毛裸蕨P. sargentii_2	张钢民 6111	四川巴塘	20010718
川西金毛裸蕨Paragymnopteris bipinnata_1	张宪春 2684	四川雅砻	20131003
川西金毛裸蕨Paragymnopteris bipinnata_2	张宪春 6942	四川冕宁	20131003
耳羽金毛裸蕨P. bipinnata var. auriculata_1	张钢民 ZGM628	陕西洋县	20230730
耳羽金毛裸蕨P. bipinnata var. auriculata_2	张钢民 436	云南丽江	20010825
耳羽金毛裸蕨P. bipinnata var. auriculata_3	张钢民 2014073	云南大理	20140814
耳羽金毛裸蕨P. bipinnata var. auriculata_4	张树仁 GGS10331	四川甘孜	20180719
金毛裸蕨P. vestita_1	罗依可 2023080460	西藏吉隆	20230804
金毛裸蕨P. vestita_2	张宪春 6441	云南丽江	20100925
金毛裸蕨P. vestita_3	张宪春7603	云南丽江	20150831
泽泻蕨Parahemionitis cordata	张宪春 3530	海南保亭	20041219
Pellaea andromedaefolia	G. DeNevers 858	美国	19810425
P. breweri	B. Bartholomew 5269	美国	19900616
P. atropurpurea	H. Koyama et al. 6633	美国	19800512
P. nana	R. Coveny & P. Hind 9476	澳大利亚	19770611
P. dura	张宪春 9065	马达加斯加	20170919
P. angulosa	张宪春 9076	马达加斯加	20170919
P. boivinii	张宪春 9197	马达加斯加	20170919
P. goudotii	张宪春 9199	马达加斯加	20170926
三角羽旱蕨P. calomelanos	张宪春9175	马达加斯加	20170924
P. striata	张宪春9198	马达加斯加	20170920
戟叶黑心蕨Calciphilopteris ludens	李保贵20010139	云南勐腊	20010417

Table 1

Fig.1

Fig.2

Fig.3

References 45

1	GASTONY G J， ROLLO D R.Phylogeny and generic circumscriptions of cheilanthoid ferns（Pteridaceae： Cheilanthoideae） inferred from rbcL nucleotide sequences［J］.American Fern Journal，1995，85（4）：341-360.
2	GASTONY G J， ROLLO D R.Cheilanthoid ferns（Pteridaceae：Cheilanthoideae） in the southwestern United States and adjacent Mexico：a molecular phylogenetic reassessment of generic lines［J］.Aliso，1998，17（2）：131-144.
3	KIRKPATRICK R E B.Investigating the monophyly of Pellaea（Pteridaceae） in the context of a phylogenetic analysis of cheilanthoid ferns［J］.Systematic Botany，2007，32（3）：504-518.
4	WINDHAM M D， HUIET L， SCHUETTPELZ E，et al.Using plastid and nuclear DNA sequences to redraw generic boundaries and demystify species complexes in cheilanthoid ferns［J］.American Fern Journal，2009，99（2）：128-132.
5	SHING K H.A new genus，Paragymnopteris Shing separated from Gymnopteris Bernh［J］.Indian Fern Journal，1993，10（1/2）：226-231.
6	ZHANG G M， YATSKIEVYCH G， HOOPER E A，et al.Subfam.Cheilanthoideae［M］//WU Z Y，RAVEN P H，HONG D Y，et al.Flora of China：Vol.2/3.Beijing：PressScience，2013：216-237.
7	DESVAUX A N.Journal de botanique，appliquée à l'agriculture，à la pharmacie，à la médicine et aux arts［M］.Paris：Bureau Du Journal De Botanique，1813.
8	CHRISTENSEN C.Index Filicum［M］.Hafniae：H.Hagerup，1906.
9	PICHI SERMOLLI R E G.Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi［J］.Webbia，1977，31（2）：313-512.
10	FRASER-JENKINS C R.New species syndrome in Indian pteridology and the ferns of Nepal［M］.India：International Book Distributors，1997.
11	FRASER-JENKINS C R， DULAWAT C S.A summary of Indian cheilanthoid ferns and the discovery of Negripteris（Pteridaceae），an Afro-Arabian fern genus new to India［J］.The Fern Gazette，2009，18（5）：216-229.
12	TRYON R M， WEATHERBY C A.A revision of the American species of Notholaena ［J］.Contributions from the Gray Herbarium of Harvard University，1956，179：1-102.
13	SMITH J.Historia Filicum：an exposition of the nature，number and organography of ferns［M］.London：Macmillan and Co.，1875.
14	TRYON R， TRYON A.Proposal to conserve Notholaena with the type Pteris trichomanoides ［J］.Taxon，1980，29（1）：160-161.
15	CHING R C.Some new nomenclatural combinations of ferns［J］.Journal of Systematics and Evolution，1965，10（4）：301-304.
16	秦仁昌，邢公侠.中国植物志：第3卷第1分册［M］.北京：科学出版社，1990.
	CHING R C， XING G X.Flora Reipublicae Popularis Sinicae：Tomus3：Part 1［M］.Beijing：Science Press，1990.
17	MICKEL J T.A redefinition of the genus Hemionitis ［J］.American Fern Journal，1974，64（1）：3-12.
18	TRYON R.Some new names and combinations in Pteridaceae［J］.American Fern Journal，1986，76（4）：184-186.
19	TRYON R， TRYON A， KRAMER K U.Pteridaceae［M］//KRAMER K U，GREEN P S.The families and genera of vascular plants：Vol 1：pteridophytes and gymnosperms.Berlin：Springer，1990：230-256.
20	YESILYURT J C， BARBARÁ T， SCHNEIDER H，et al.Identifying the generic limits of the cheilanthoid genus Doryopteris ［J］.Phytotaxa，2015，221（2）：101-122.
21	PPG I.A community-derived classification for extant lycophytes and ferns［J］.Journal of Systematics and Evolution，2016，54（6）：563-603.
22	FIELD A R.Classification and typification of Australian lycophytes and ferns based on Pteridophyte Phylogeny Group classification PPG I［J］.Australian Systematic Botany，2020，33（1）：1-102.
23	WANG W， YANG W L， MAO X X，et al.The phylogenetic affinities of Pellaea connectens，a rare endemic Chinese fern［J］.Phytotaxa，2015，220（1）：30-42.
24	ZHANG G M， ZHANG X C， CHEN Z D，et al.First insights in the phylogeny of Asian cheilanthoid ferns based on sequences of two chloroplast markers［J］.Taxon，2007，56（2）：369-378.
25	SCHUETTPELZ E， SCHNEIDER H， HUIET L，et al.A molecular phylogeny of the fern family Pteridaceae：assessing overall relationships and the affinities of previously unsampled genera［J］.Molecular Phylogenetics and Evolution，2007，44（3）：1172-1185.
26	ROTHFELS C J， WINDHAM M D， GRUSZ A L，et al.Toward a monophyletic Notholaena（Pteridaceae）：resolving patterns of evolutionary convergence in xeric‐adapted ferns［J］.Taxon，2008，57（3）：712-724.
27	LIU H M.Embracing the pteridophyte classification of Ren‐Chang Ching using a generic phylogeny of Chinese ferns and lycophytes［J］.Journal of Systematics and Evolution，2016，54（4）：307-335.
28	BROWNSEY P J， OHLSEN D J， SHEPHERD L D，et al.A review of the fern genus Pellaea（Pteridaceae） in Australasia［J］.Australian Systematic Botany，2020，33（5）：446-457.
29	GRUSZ A L， WINDHAM M D， PICARD K T，et al.A drought-driven model for the evolution of obligate apomixis in ferns：evidence from pellaeids（Pteridaceae）［J］.American Journal of Botany，2021，108（2）：263-283.
30	PRADO J， DEL NERO RODRIGUES C， SALATINO A，et al.Phylogenetic relationships among Pteridaceae，including Brazilian species，inferred from rbcL sequences［J］.Taxon，2007，56（2）：355-368.
31	SÄRKINEN T， STAATS M， RICHARDSON J E，et al.How to open the treasure chest？ Optimising DNA extraction from herbarium specimens［J］.PLoS One，2012，7（8）：e43808.
32	KATOH K， MISAWA K， KUMA K I，et al.MAFFT：a novel method for rapid multiple sequence alignment based on fast Fourier transform［J］.Nucleic Acids Research，2002，30（14）：3059-3066.
33	KATOH K， STANDLEY D M.MAFFT multiple sequence alignment software version 7：improvements in performance and usability［J］.Molecular Biology and Evolution，2013，30（4）：772-780.
34	BOLGER A M， LOHSE M， USADEL B.Trimmomatic：a flexible trimmer for Illumina sequence data［J］.Bioinformatics，2014，30（15）：2114-2120.
35	JIN J J， YU W B， YANG J B，et al.GetOrganelle：a fast and versatile toolkit for accurate de novo assembly of organelle genomes［J］.Genome Biology，2020，21：241.
36	FRANCHET M A.Plantae davidianae ex sinarum im- perio［J］.Nouvelles archives du muséum d'histoire naturelle，1887，2（10）：123.
37	CHING R C.New and little known ferns from China and neighbor［J］.Lingnan Science Journal，1936，15（3）：398-399.
38	秦仁昌.中国蕨类植物小志［J］.植物分类学报，1982，20（2）：233.
	CHING R C.Notes on some Chinese ferns［J］.Journal of Systematics and Evolution，1982，20（2）：233.
39	张宪春.中国石松类和蕨类植物［M］.北京：北京大学出版社，2012.
	ZHANG X C.Lycophytes and ferns of China［M］.Beijing：Peking University Press，2012.
40	LINNAEUS C， SALVIUS L.Species plantarum：Vol.2［M］.Holmiae：Impensis Laurentii Salvii，1753：1071.
41	JARVIS C.Order out of chaos：linnaean plant names and their types［M］.London：Linnean Society of London in Association with the Natural History Museum，2007：252-342.
42	BAKER J G.A summary of the new ferns which have been discovered or described since 1874［J］.Annals of Botany，1891，5（19）：301-332.
43	HOOKER W J.Icones plantarum：Vol.2［M］.London：Longman，Rees，Orme，Brown，Green，& Longman，1837：115.
44	CHRIST H.Filices Chinae Occidentalis［J］.Bulletin de l'Académie Internationale de Géographie Botanique，1906，16（199/200/201）：130.
45	CHRIST H.Filices Wilsonianae［J］.Botanical Gazette，1911，51（5）：345-359.

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