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Classification, Molecular Phylogeny, Divergence Time, And The JapaneseSocietyJapanese Society for Plant Systematics ISSN 1346-7565 Acta Phytotax. Geobot. 56 (2): 111-126 (2005) Invited article and Classification,MolecularPhylogeny,DivergenceTime, Morphological Evolution of Pteridophytes with Notes on Heterospory and and Monophyletic ParaphyleticGroups MASAHIRO KATO* Department ofBiotogicat Sciences,Graduate Schoot ofScience,Universitv. of7bkyo, Hongo, 7bk)]o IJ3- O033, lapan Pteridophytes are free-sporing vascular land plants that evolutionarily link bryophytes and seed plants. Conventiona], group (taxon)-based hierarchic classifications ofptcridophytes using phenetic characters are briefiy reviewcd. Review is also made for recent trcc-based cladistic analyses and molecular phy- logenetic analyses with increasingly large data sets ofmultiplc genes (compared to single genes in pre- vious studies) and increasingly large numbers of spccies representing major groups of pteridophytes (compared to particular groups in previous studies), and it is cxtended to most recent analyses of esti- mating divergcnce times ofpteridephytes, These c]assifications, phylogenetics, and divergcncc time esti- mates have improved our understanding of the diversity and historical structure of pteridophytes. Heterospory is noted with referencc to its origins, endospory, fertilization, and dispersal. Finally, menophylctic and paraphyletic groups rccently proposed or re-recognized are briefly dcscribcd. Key words: classification, divergence timc estimate. fems,heterospory, molecular phylogcny, pteri- dophytcs. Morphological Classifications it a recent diversification, resulting in a total of about 12,OOO species, which may be primitive or Pteridophytes, like seed plants (gymnosperms and advanced. The species were classified in many difl angiosperms), are vascular land plants and also are ferent classificatien systems based on morphologi- similar to nonvascular bryophytes in the free-speringcal characters. Some of major classifications put reproduction, Evolutionarily they fo11owed bryo- fbrward in the 20th century are briefly noted here phytes and preceded seed plants. Thus, free-sporing ('lables 1, 2). vascular plants or pteridophytes in a broad sense Engler & Prantl (1902) classified pteridophytes have a long (420 million years) evolutionary histo- into fbur classes, Filicales, Sphenophyllales, ry, and on the other hand, like angiosperms, exhib- Equisetales and fycopodia]es, and subdivided the * PTesent address: Department of Botany, National Science Museum, Tsukuba 305-OO05, Japan e-mail address, [email protected] This article is fonned frem the presentation as one of contributions for the International Symposium 2004, Asian Plant Diversity and Systematics, held at Sakura, Chiba, Japan on July 29 - August 2, 2004, NII-Electronic Library Service The JapaneseSocietyJapanese Society for PlantPlantSystematics Systematics 112 APG Xlo1. 56 TABLE 1 . Classifications ofmajoT ptcridophytc groups proposed by sorne authors, based on cemparative morphology, Numbers indi- cate grouping in each classification and do not correspond among classifications. Engler & Prantt C1902) Verdooun (1938)Tagawa&Iwatsuki(1972)PichiSermolli(1977)Chmg{E97S)Tryon & TT)'on (1982) Krarner & Green (1990) 1,Lycopodiales 1,Lycopodiinae 1.Lycopsida 1.Lycephytina 1,Lycephytina1.Lycoodiepsida 1.Lycopodiatae 1-1.Ligulatae 1-1-t.Se]aginellineae1-1.Selaginel]ales1-1.Se]agine]]ales 1-1.Selaginel[ales1-1.SeiaginellRlesl-1,Selaginellales1-1,Selaginellales 1-I-2.Isoetineae l-2.Isoetales 1-2.Isoetales 1-2,Jsoetales 1-2. Lycopodia]cs ]-2, Isoctales 1-2,lsoetales 1-2.Eligulatae 1-2-1.LycopDdincac 1-3.Lycopodia]es1-3.Lycopodiales 1-3.Lycopodjales 2.Isocphytina 1-3.Lycopodiales1-3,Lycepodiales 1-2-2.Psilotineae 2.Psi]ophytinae 2,Psi]opsida 2,?sjophytina 3,Psjophytina 2.Psilotatae 2.Equiestales 3.Articulatae 3,Equisetopsida 3.Sphenophytina4, Sphenophytina 2.Equisetopsida 3.Equisetatae 3,SphenophylLates 4.Filica]es 4. Filicinae 4.Pteropsida 4.Filicophytina 5.Fi]icophytina3.Fiticopsida 4.Filicatae (incl.Psilotaceae) Extinct group included in Equisetopsida in other classifications, TABi.E 2. Classifications ofmajor fern groups proposed by some authors, based on comparative morphology. Numbers indicate group- ing in each classification and do not correspond among classifications. Christellsen(l938)Cepeland(1947)Tagawa&iwatsuki(1972)PichiSermolli(1977)Ching{IY7S) Tryon&Tryen(1982) 1,Eusporangiatae 1.Eusporangiopsida 1,Polypodiidae 1-].Ophioglossales ] . Qphioglossales1.0phioglossales 1,Ophioglossopsida 1-1,Ophioglossales 1-1.0phioglossales 1-2,Maradiaies 2.Marattiales 2.Maraniales 2.Marattiepsida 1-2.Marattia]es 1-2.Marattiales 2,Leptosporangiatae3. FiHcales 3.Filicales 3,Fllicopsida 2,Protoleptosporangiopsida1-3.PojypodiaLes 2-1.Filicales 3-].Osmundjclae 3.Leptosporangiopsida t-3-1.Po]ypodiineae 3-2,Plagiogyriidae 3-t,Po]ypodiales 3-3.Gleicheniidae 3-4.Schizaeidae 3-5,Hymenophyllidae 2-2, Salviniales 4. Salviniales 3-6,Salvinlidae 3-2,Salviniales t-3-2.Salviniineae 5.Marsileales 3-1,MarsiTeidae 3-3,Marslleales 1-3-3.Marsiteineae 2. Psilotidae class Filicales into three orders, Filicales leptospo- and in total 14 families including the Iarge family rangiatae (suborders Eufilicineae and Hydro- Polypodiaceae with 15 subfamilies. Ching (1940) "Polypodiaceae" pteridineae), Maiattiales and Ophioglossales, and the classified into 33 families and rec- "Poly- class Lycopodiales into two orders Lycopodiales ognized five series in the polyphyletic eligulatae including suborder Psilotineae and podiaceae" in the context of phylogeny, Ching Lycepodiales ligulatae with suborders Selaginetli-(1978) classified Chinese pteridophytes (division neae and Isoetineae, Christensen's (1905, 1913- Pteridophyta) into five subdivisions: subdiv, 1934) Index Filicum and supplements I-III enu- Lycophytina comprising orders Lycopodiales and merated all fern species ofthe world described, In SeJaginellales, three monotypic subdiv, Isoephytina, his systematic classification of ferns Christensen Sphenophytina and Psilophytina, and subdiv. (1938) recognized two series (Filices EusporangiataeFilicophytina comprising three classes (Eusporan- and Filices Leptosporangiatae), two orders giopsida [orders Ophioglossales and Marattiales], Ophioglossales and Marattiales in the former series Protoleptosporangiopsida [Osmundales], and and two orders Filicales and Salviniales in the latteg Leptosporangiopsida [Polypodiales or Filicales, NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics August 2005 KPLTO: PteridophyLe phylogeny and evelution 113 Marsileales, Salviniales]). In Genera Filicum (suborders Polypodiineae, Marsileineae, and Copeland (1 947) recognized orders Ophioglossales Salviniineae), and divided Lycopodiopsida into (1 family), Marattiales (1 family), and Filicales (19 bycopodiales, Selaginellales and Isoetalcs. Ttyon & families including Marsileaceae and Salviniaceae), Tryon (1982) included Psilotaceae in Filicopsida Holttum (1949) classified leptosporangiate ferns (ferns) based on BierhorsVs (1977 and references into 14 families including the large family Denn- cited therein) morphological and anatomical results staedtiaceae with 1 1 subfamilies and proposed three and spore wall characters. phylegenetic lineages, ofwhich one is terrninated by In short, the classifications based on phenetic rlagawa Dennstaedtiaceae. & Iwatsuki (1972) adopt- characters usually recognized four major groups ed the conventional classification ofpteridophytes of living pteridophytes, among which ferns were into four classes Psiopsida, Lycopsida, Equiset- divided into Marattiales, Ophioglossales and opsida, and Pteropsida. They classjfied Pteropsida Filicales, although certain groups (e.g., Psilotaceae into erders Ophioglossales, Marattiales, Filicalcs, and aquatic ferns) were assigned to different groups Marsileales, and Salviniales. [lagawa & Iwatsuki ofhigher ranks or treated at different ranks ([lables (1972) recognized in total 34 families for pterido- 1, 2). Characters that are infbrmative throughout phytes of Thailand. Pichi Sermolli (1977) classified pteridophytes are not many. Those classifications Pteridophyta into fbur subdivisions, Lycophytina, with hierarchic ranks are generally taxon-based and Sphenophytina, Psilophytina, and Filicophytina, have usually not been given statistically analyzed The first three were monotypic each with single interrelationships of families. classes, while the last Filicophytina comprised three classes, Ophioglossopsida, Marattiopsida and Molecular Phylogenies Filicopsida. Pichi Sermolli (1977) assigned 58 of64 families to Filicopsida, 3 to Lycopsida, 1 to Achievement ofmelecular phylogeny, which is dis- Equisetopsida, and 2 to Psilotopsida. Kramer & played as a phylegenetic tree, succeeded long con- Green (1990) compiled contributions to pterido- tributions of systematics, classifications, and fio- phyte classification and presented a similar classi- ras based on phenetic or morphological characters, fication system of four classes Psilotatae (1 family), as noted above, Molecular analyses with large data Lycopodiatae (3 families), Equisetatae (1 family), sets dealing with all or most pteridophyte groups and Filicatae (33 families). Among families of have been explosive since the middle 1990s. One Filicatae, affinities were suggested between year later than Chase et al.'s (1993) epoch-making Dipteridaceae and Cheiropleuriaceae; Vittariaceaestudy on angiosperm phylogeny using a large data and Pteridaceae; tree fern families; Lomariopsida-
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