Stasis and convergence characterize morphological evolution in eupolypod II ferns

Annals of Botany

Volumn 113, Issue 1, 2014, Pages 35-54

  Sundue, Michael A ^a   Rothfels, Carl J ^b  

^a UNIVERSITY OF VERMONT   (United States)

^b DUKE UNIVERSITY   (United States)

Author keywords

Athyriaceae; character state reconstruction; convergent evolution; Diplaziopsidaceae; eupolypods II; morphological evolution; Polypodiales; rate of evolution; Rhachidosoraceae; symplesiomorphy; Woodsiaceae

Indexed keywords

CHROMOSOME; CLADISTICS; EVOLUTION; FERN; MORPHOLOGY; PARSIMONY ANALYSIS; PHYLOGENY; POPULATION GENETICS; VASCULAR PLANT;

ARTICLE; ATHYRIACEAE; BAYES THEOREM; CHARACTER STATE RECONSTRUCTION; CONVERGENT EVOLUTION; DIPLAZIOPSIDACEAE; EUPOLYPODS II; EVOLUTION; FERN; HISTOLOGY; MORPHOLOGICAL EVOLUTION; PHYLOGENY; PLESIOMORPHY; POLYPODIALES; RATE OF EVOLUTION; RHACHIDOSORACEAE; STATISTICAL MODEL; WOODSIACEAE;

ATHYRIACEAE; CHARACTER STATE RECONSTRUCTION; CONVERGENT EVOLUTION; DIPLAZIOPSIDACEAE; EUPOLYPODS II; MORPHOLOGICAL EVOLUTION; POLYPODIALES; RATE OF EVOLUTION; RHACHIDOSORACEAE; SYMPLESIOMORPHY; WOODSIACEAE;

BAYES THEOREM; BIOLOGICAL EVOLUTION; FERNS; LIKELIHOOD FUNCTIONS; PHYLOGENY;

EID: 84891505997     PISSN: 03057364     EISSN: 10958290     Source Type: Journal    
DOI: 10.1093/aob/mct247     Document Type: Article

References (178)

1. Atkinson LR. 1973. The gametophyte and family relationships. In: Jermy AC, Crabbe JA, Thomas BA, eds. The phylogeny and classification of ferns. London: Linnean Society, 73-90
2. Atkinson LR, Stokey AG. 1964. Comparative morphology of the gametophyte of homosporous ferns. Phytomorphology 14: 51-70
3. Bir SS. 1969a. Observations on the morphology and anatomy of Diplaziopsis javanica (Bl) C. Ch. Plant Science 1: 110-118
4. Bir SS. 1969b. The stelar anatomy of Diplazium latifolium Moore. American Fern Journal 59: 23-26
5. Bir SS. 1971. Anatomy of some Himalayan species of Athyrium. Journal of the Indian Botanical Society 50: 247-252
6. Bir SS, Trikha CK. 1970. Anatomical observations on the genus Cystopteris Bernhardi. Proceedings of the Indian National Science Academy 36B: 192-204
7. Blasdell RF. 1963.A monographic study of the fern genus Cystopteris. Memoirs of the Torrey Botanical Club 21: 1-102
8. Bosman MTM. 1991. A monograph of the fern genus Microsorum (Polypodiaceae). Leiden Botanical Series 14: 1-161
9. Bower FO. 1914. Studies in the phylogeny of the filicales, IV. Blechnum and allied genera. Annals of Botany 28: 363-343
10. Bower FO. 1923. The Ferns, Treated Comparatively with a View to their Natural Classification. Cambridge: Cambridge University Press
11. Bower FO. 1928. The ferns, treated comparatively with a view to their natural classification. Cambridge: Cambridge University Press
12. Brown DFM. 1964.A monographic study of the fern genusWoodsia Beih. Nova Hedwigia 16: 1-156
13. Brown R. 1810. Prodromus Florae Novae Holandiae et Insulae Van-Diemen. London: Richard Taylor & Sons
14. Campbell DH. 1905. The structure and development of mosses and ferns. London: MacMillan
15. Campbell DH. 1940. The evolution of the land plants (Embryophyta). Stanford: Stanford University Press
16. Chandra S, Nayar BK. 1970. Some aspects of the morphology of the rhizome of Pteris wallichiana. Proceedings: Plant Sciences 71: 79-85
17. Ching R-C. 1936. A revision of the Chinese and Sikkim-Himalayan Dryopteris with reference to some species from neighbouring regions. Bulletin of the Fan Memorial Institute of Biology 6: 237-352
18. Ching R-C. 1940. On natural classification of the familyPolypodiaceaé. Sunyatsenia 5: 1-37
19. Ching R-C. 1964. On some confused genera of the family Athyriaceae. Acta Phytotaxonomica Sinica 9: 41-84
20. Ching R-C. 1978. The Chinese fern families and genera: Systematic arrangement and historical origin. Acta Phytotaxonomica Sinica 16: 1-19
21. Christensen C. 1913. A monograph of the genus Dryopteris part I, the tropical American pinnatifid-bipinnatifid species. Skrifter Udgivne af Videnskabs-Selskabet i Christiana. Mathematisk-naturvidenskabelig Klass 7.x.2
22. Christensen C. 1938. Filicinae. In: Verdoorn F, ed. Manual of pteridology. The Hague: Martinus Nijhoff
23. Copeland EB. 1947. Genera Filicum.Waltham: Chronica Botanica Co
24. Cranfill R, KatoM.2003. Phylogenetics, biogeography, and classification of the woodwardioid ferns (Blechnaceae). In: Chandra S, Srivastava M, eds. Pteridology in the new millennium. Dordrecht: Kluwer Academic Publishers
25. Crowden RK, Jarman SJ. 1974. 3-deoxyanthocyanins from the fern Blechnum procerum. Phytochemistry 13: 1947-1948
26. Davies KL. 1991. A brief comparative survey of aerophore structure within the Filicopsida. Botanical Journal of the Linnean Society 107: 115-137
27. Diels FLE. 1899. Woodsieae in Engler H.G.A., Prantl K.A.E., Naturlichen Pflanzenfamilien I, 4: 158-159
28. Freckleton RP, Harvey PH, Pagel MD. 2002. Phylogenetic analysis and comparative data: A test and review of evidence. American Naturalist 160: 712-726
29. Gantt E, Arnott HJ. 1965. Spore germination and development of the young gametophyte of the ostrich fern (Matteuccia struthiopteris). American Journal of Botany 52: 82-94
30. Goebel K. 1900. Organography of plants, especially of the Archegoniata and the Spermatophyta. Oxford: Clarendon Press
31. Goloboff PA, Farris JS, Nixon KC. 2008.TNT, a free program for phylogenetic analysis. Cladistics 24: 774-786
32. Harmon LJ, Weir JT, Brock CD, Glor RE, Challenger W. 2007. GEIGER: Investigating evolutionary radiations. Bioinformatics 24: 129-131
33. Hasebe M, Wolf PG, Pryer KM, et al. 1995. Fern phylogeny based on rbcL nucleotide sequences. American Fern Journal 85: 134-181
34. Hayata B. 1927. On the systematic importance of the stelar system in the Filicales. Bot. Mag. (Tokyo) 41: 697-718
35. Hennipman E, Roos MC. 1982. A monograph of the fern genus Platycerium (Polypodiaceae). Amsterdam: North-Holland Publishing Co
36. Hernández-Hernández V, Terrazas T, Mehltreter K, Angeles G. 2012. Studies of petiolar anatomy in ferns: Structural diversity and systematic significance of the circumendodermal band. Botanical Journal of the Linnean Society 169: 596-610
37. Holttum RE. 1940. The fern genus Diplazium in the Malay Peninsula. Gardens Bulletin Straights Settlement 11: 74-108
38. Holttum RE. 1947. A revised classification of leptosporangiate ferns. The Journal of the Linnean Society of London. Botany 53: 123-155
39. HolttumRE. 1949. The classification of ferns. BiologicalReviews 24: 267-296
40. Holttum RE. 1954. A revised flora of Malaya. Singapore: Government Printing Office
41. Holttum RE. 1958. Notes on Malaysian ferns, with descriptions of a new genus and new species. Kew Bulletin 13: 447-455
42. Holttum RE. 1959. Some notes on vegetative characters of ferns. Brittish Fern Gazette 9: 2-4
43. Holttum RE. 1960. Vegetative characters distinguishing the various groups of ferns included in Dryopteris of Christenseńs Index Filicum, and other ferns of similar habit and sori. Gardenś Bulletin, Singapore 17: 361-367
44. Holttum RE. 1982. Thelypteridaceae. In: Van Steenis CGGJ, Holttum RE, eds. 1959-1982. Flora Malesiana. Series II. Pteridophyta. Vol. 1, part 5
45. Hovenkamp P. 1986. A monograph of the fern genus Pyrrosia (Polypodiaceae). Leiden: E.J. Brill/Leiden University Press
46. Hovenkamp P. 1996. The inevitable instability of generic circumscriptions in Old World Polypodiaceae. In: Camus JM, Gibby M, Johns RJ, eds. Pteridology in perspective. Kew: Royal Botanical Gardens
47. Huang T-C. 1981. Spore flora of Taiwan. Taiwan: Tah-Jinn Press Co/National Taiwan University
48. Iwatsuki K. 1970. Taxonomic studies of Pteriodophyta IX. Acta Phytotaxonomica et Geobotanica 4-6: 182-188
49. Janssen T, Schneider H. 2005. Exploring the evolution of humus collecting leaves in drynarioid ferns (Polypodiaceae, Polypodiidae) based on phylogenetic evidence. Plant Systematics and Evolution 252: 175-197
50. Jian S, Soltis PS, Gitzendanner MA, et al. 2008. Resolving an ancient, rapid radiation in Saxifragales. Systematic Biology 57: 38-57
51. Johnson AK, Rothfels CJ, Windham MD, Pryer KM. 2012. Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae). American Journal of Botany 99: 1118-1124
52. Johnson DM. 1986. Trophopods in North American species of Athyrium (Aspleniaceae). Systematic Botany 11: 26-31
53. Kato M. 1972. The vascular structure and its taxonomic significance in the Athyriaceae. Acta Phytotaxonomica et Geobotanica 25: 79-91
54. Kato M. 1973. Taxonomical evaluation of the articulated hairs found in the Athyriaceae. Acta Phytotaxonomica et Geobotanica XXV: 119-126
55. Kato M. 1975. On the systematic position of Athyrium mesosorum. Acta Phytotaxonomica et Geobotanica 27: 56-60
56. Kato M. 1977. Classification of Athyrium and allied genera of Japan. Botanical Magazine (Tokyo) 90: 23-40
57. Kato M. 1979a. A new combination in Athyrium (Athyriaceae) of North America. Annals of Carnegie Museum 49: 177-179
58. Kato M. 1979b. Taxonomic study of the genus Cornopteris (Athyriaceae). Acta Phytotaxonomica et Geobotanica 30: 101-118
59. Kato M. 1984.Ataxonomic study of the athyrioid fern genus Deparia with main reference to the Pacific species. Journal of theFaculty of Science, University of Tokyo, Section III. Botany 13: 375-430
60. Kato M, Darnaedi D. 1988. Taxonomic and phytogeographic relationships of Diplazium flavoviride, D. pycnocarpon, and Diplaziopsis. American Fern Journal 78: 77-85
61. Kato M, Kramer K. 1990. Athyrioideae. In: Kramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer
62. Keating RC. 1968.Trends of specialization in the stipe anatomyof Dennstaedtia and related genera. American Fern Journal 58: 126-140
63. Kenrick P, Crane P. 1997. The origin and early diversification of land plants: A cladistic study.Washington, DC: Smithsonian Institution Press
64. Kirkpatrick REB. 2007. Investigating the monophyly of Pellaea (Pteridaceae) in the context of a phylogenetic analysis of cheilanthoid ferns. Systematic Botany 32: 504-518
65. Knapp R. 2011. Ferns and fern allies of Taiwan. Taiwan: KBCC Press
66. Korall P, Conant DS, Schneider H, Ueda K, Nishida H, Pryer KM. 2006. On the phylogenetic position of Cystodium: It́s not a tree fern-It́s a polypod! American Fern Journal 96: 45-53
67. Kramer KU. 1987. A brief survey of the dromy in fern leaves, with an expanded terminology. Botanica Helvetica 97: 219-228
68. Kramer KU, Green PS. 1990. Pteridophytes and gymnosperms. InKramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer-Verlag
69. Kramer KU, Viane R. 1990. Aspleniaceae. In: Kramer KU, Green PS eds. The families and genera of vascular plants. Berlin: Springer-Verlag
70. Kramer KU, Chambers TC, Hennipman E. 1990a. Blechnaceae. In: Kramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer-Verlag
71. Kramer KU, Holttum RE, Moran RC, Smith AR. 1990b. Dryopteridaceae. In: Kramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer-Verlag
72. Kreier H-P, Schneider H. 2006. Phylogeny and biogeography of the staghorn fern genus Platycerium (Polypodiaceae, Polypodiidae). American Journal of Botany 93: 217-225
73. Kuo L-Y, Li F-W, Chiou W-L, Wang C-N. 2011. First insights into fern matK phylogeny. Molecular Phylogenetics and Evolution 59: 556-566
74. Labiak PH, Rouhan G, Sundue MA. 2010a. Phylogeny and taxonomy of Leucotrichum (Polypodiaceae): A new genus of grammitid ferns from the Neotropics. Taxon 59: 911-921
75. Labiak PH, Sundue MA, Rouhan G. 2010b. Molecular phylogeny, character evolution, and biogeography of the grammitid fern genus Lellingeria (Polypodiaceae). American Journal of Botany 97: 1354-1364
76. Lellinger DB. 2002.Amodern multilingual glossary for taxonomic pteridology. Pteridologia 3: 1-263
77. Lewis PO. 2001. A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology 50: 913-925
78. Liew FS. 1971. Studies of North American Athyrium. New York: Columbia University
79. Lin B-L, DeVol CE. 1977. The use of stipe characters in fern taxonomy I. Taiwania 22: 91-99
80. Lin B-L, DeVol CE. 1978. The use of stipe characters in fern taxonomy II. Taiwania 23: 77-95
81. Linnaeus C. 1753. Species plantarum: Exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Stockholm: Impensis Laurentii Salvii
82. Liu Y-C, Chiou W-L, Liu H-Y. 2009. Athyrium. In. Flora of Taiwan. Taipei, Taiwan: Taiwan Forestry Research Institute
83. Lloyd RM. 1971. Systematics of the onocleoid ferns. University of California Publications in Botany 61: 1-87
84. Lloyd RM, Klekowski EJr. 1970. Spore germination and viability in Pteridophyta: Evolutionary significance of chlorophyllous spores. Biotropica 2: 129-137
85. Maddison W, Maddison D. 2006. Mesquite:Amodular system for evolutionary analysis. mesquiteproject.org
86. Manton I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge: Cambridge University Press
87. Manton I, SledgeW. 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Philosophical Transactions B 238: 127-184
88. MarengoNP. 1949.Astudy of the cytoplasmic inclusions during sporogenesis in Onoclea sensibilis. American Journal of Botany 36: 603-613
89. MasuyamaS. 1975a. The sequence of the gametangium formation in homosporous fern gametophytes II. Types and their taxonomic distribution. The Science Reports of the Tokyo Kyoiku Daigaku Sec. B 16: 71-86
90. MasuyamaS. 1975b. The sequence of the gametangium formation in homosporous fern gametophytes. I. Patterns and the possible effect on the fertilization, with special reference to the gametophytes of Athyrium. The Science Reports of the Tokyo Kyoiku Daigaku Sec. B. 16: 47-69
91. Mendoza-Ruiz A, Pérez-García B. 2009. Morphogenesis of the gametophytes of eight Mexican species of Blechnum (Blechnaceae). Acta Botanica Mexicana 88: 59-72
92. Mickel JT. 2003. Ferns for American gardens. Portland, OR: Timber Press
93. Mickel JT, Smith AR. 2004. The pteridophytes of Mexico. New York: The New York Botanical Garden Press
94. Momose S. 1958. Prothalia of onocleoid ferns. The Journal of Japanese Botany 33: 289-296
95. Momose S. 1959. The prothalia of Aspleniaceae (2). The Journal of Japanese Botany 34: 293-302
96. Momose S. 1960a. The prothalia of Aspleniaceae (3). The Journal of Japanese Botany 35: 47-54
97. Momose S. 1960b. The prothalia of Aspleniaceae (4). The Journal of Japanese Botany 35: 299-235
98. Momose S. 1964. The prothalium of Cryptogramma group of ferns. The Journal of Japanese Botany 39: 305-312
99. Momose S. 1965a. Prothalia of some ferns from the Galapagos Islands (1). The Journal of Japanese Botany 40: 140-147
100. Momose S. 1965b. Prothalia of some ferns from the Galapagos Islands (2). The Journal of Japanese Botany 40: 277-281
101. Moran RC. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural History Survey Bulletin 34
102. Moran RC. 1991. Monograph of the Neotropical fern genus Stigmatopteris (Dryopteridaceae). Annals of the Missouri Botanical Garden 78: 857-914
103. Moran RC, Prado J. 2010. Megalastrum (Dryopteridaceae) inCentral America. Kew Bulletin 65: 137-188
104. Moran RC, Hanks JG, Rouhan G. 2007. Spore morphology in relation to phylogeny in the fern genus Elaphoglossum(Dryopteridaceae). International Journal of Plant Sciences 168: 905-929
105. Moran RC, Hanks JG, Labiak P, Sundue M. 2010a. Perispore morphology of bolbitidoid ferns (Dryopteridaceae) in relation to phylogeny. International Journal of Plant Science 171: 872-881
106. Moran RC, Labiak PH, Sundue M. 2010b. Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia 62: 337-356
107. MortonCV. 1959. The identification of aCosta Rican Blechnum. AmericanFern Journal 49: 66-69
108. MortonCV. 1963. The classification of Thelypteris. American Fern Journal 53: 149-154
109. Murakami N, Moran RC. 1993. Monograph of the neotropical species of Asplenium sect. Hymenasplenium (Aspleniaceae). Annals of the Missouri Botanical Garden 80: 1-38
110. Nakazato T, Barker MS, Rieseberg LH, Gastony GJ. 2008. Evolution of the nuclear genome of ferns and lycophytes. In: Ranker TA, Haufler CH, eds. Biology and evolution of ferns and lycophytes. Cambridge: Cambridge University Press
111. Nauman CE. 1989. Gametophyte development and morphology in seven neotropical taxa of Nephrolepis Schott. Phytomorphology 39: 29-39
112. Nayar BK, Bajpai N, Raza F. 1966. Morphological studies on some species of Blechnum, Doodia, Woodwardia and Stenochlaena: I. The gametophytes and the juvenile sporophytes. Journal of the Linnean Society of London. Botany 59: 405-423
113. Nayar BK, Kaur S. 1971. Gametophytes of homosporous ferns. The Botanical Review 37: 295-396
114. Nixon KC. 1999. The parsimony ratchet, a newmethod for rapid parsimony analysis. Cladistics 15: 407-414
115. Nixon KC. 1999-2002.Winclada. Ithaca, New York: Software and documentation published by the author
116. Ogura Y. 1972. Comparative anatomy of vegetative organs of the pteridophytes. Berlin: Gebruder Borntraeger
117. Pacheco L, Moran RC. 1999. Monograph of the neotropical species of Callipteris with anastomosing veins (Woodsiaceae). Brittonia 51: 343-388
118. Pagel M. 1999. Inferring the historical patterns of biological evolution. Nature 401: 877-884
119. PalmerDD.2002.Hawaiís fern and fern allies. Honolulu: University ofHawaíi Press
120. Paris CA, Barrington DS. 1990.William Jackson Hooker and the generic classification of ferns. Annals of the Missouri Botanical Garden 77: 228-238
121. Pickett FL, Thayer LA. 1927. The gametophytic development of certain ferns: Polypodium vulgare var. occidentale and Pellaea densa. Bulletin of the Torrey Botanical Club 54: 249-255
122. Prado J, Rodrigues CDN, Salatino A, SalatinoMLF. 2007. Phylogenetic relationships among Pteridaceae, including Brazilian species, inferred from rbcL sequences. Taxon 56: 355-368
123. Presl KB. 1848 (1847). Die gefässbündel im stipes der farrn: Druck der k. k. Hofbuchdr. von G. Haase Söhne
124. Price MG. 1990. Philippine fern notes. Contr. Univ. Mich. Herb. 17: 267-278
125. Pryer KM, Smith AR, Skog JE. 1995. Phylogenetic relationships of extant ferns based on evidence from morphology and rbcL sequences. American Fern Journal 85: 205-282
126. Pryer KM, Schneider H, Smith AR, et al. 2001. Horsetails and ferns are amonophyletic group and the closest living relatives to seed plants. Nature 409: 618-622
127. Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R. 2004. Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. American Journal of Botany 91: 1582-1598
128. Pryer KM, Smith AR, Rothfels CJ. 2008. Polypodiidae Cronquist, Takht. & Zimmerm. 1966. Leptosporangiate Ferns. In The Tree of Life Web Project, http://tolweb.org/
129. RDevelopmentCoreTeam. 2011. R:Alanguage and environment for statistical computing. Vienna: R Foundation for Statistical Computing
130. Ranker TA, Smith AR, Parris BS, et al. 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): A case of rampant morphological homoplasy. Taxon 53: 415-428
131. Regalado GL, Prada C. 2011. The genus Hymenasplenium (Aspleniaceae) in Cuba, including new combinations for the Neotropical species. American Fern Journal 101: 265-281
132. Rödl-Linder G. 1990. Monograph of the fern genus Goniophlebium. Blumea 34: 277-423
133. Rothfels CJ, Windham MD, Grusz AL, Gastony GJ, Pryer KM. 2008. Toward a monophyletic Notholaena (Pteridaceae): Resolving patterns of evolutionary convergence in xeric-Adapted ferns. Taxon 57: 712-724
134. Rothfels CJ, Larsson A, Kuo L-Y, Korall P, Chiou W-L, Pryer KM. 2012a. Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Systematic Biology 61: 490-509
135. Rothfels CJ, Sundue MA, Kuo L-Y, et al. 2012b. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61: 515-533
136. Sano R, Takamiya M, Ito M, Kurita S, Hasebe M. 2000a. Phylogeny of the lady fern group, tribe Physematieae (Dryopteridaceae), based on chloroplast rbcL gene sequences. Molecular Phylogenetics andEvolution 15: 403-413
137. Sano R, Takamiya M, Kurita S, Ito M, Hasebe M. 2000b. Diplazium subsinuatum and Di. tomitaroanum should be moved to Deparia according to molecular, morphological, and cytological characters. Journal of Plant Research 113: 157-163
138. Schneider H. 2007. Plant morphology as the cornerstone to the integration of fossils and extant taxa in phylogenetic analyses. Species, Phylogeny and Evolution 1: 65-71
139. Schneider H, Pryer KM, Cranfill R, Smith AR, Wolf PG. 2002. Evolution of vascular plant body plans: A phylogenetic perspective. In: Cronk QCB, Bateman RM, Hawkins JA, eds. Developmental genetics and plant evolution. London: Taylor & Francis
140. Schneider H, Russell S, Cox C, et al. 2004a. Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its implications for biogeography. Systematic Botany 29: 260-274
141. Schneider H, Schuettpelz E, Pryer KM, Cranfill R, Magallon S, Lupia R. 2004b. Ferns diversified in the shadow of angiosperms. Nature 428: 553-557
142. Schneider H, Smith AR, Pryer KM. 2009. Is morphology really at odds with molecules in estimating fern phylogeny? Systematic Botany 34: 455-475
143. Schuettpelz E, Pryer KM. 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037-1050
144. Schuettpelz E, Pryer KM. 2008. Fern phylogeny. In: Ranker TA, Haufler CH, eds. Biology and evolution of ferns and lycophytes. New York: Cambridge University Press
145. Schuettpelz E, Korall P, Pryer KM. 2006. Plastid atpA data provide improved support for deep relationships among ferns. Taxon 55: 897-906
146. Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM.2007.Amolecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera. Molecular Phylogenetics and Evolution 44: 1172-1185
147. Sheffield E, Bell PR. 1987. Current studies of the pteridophyte life cycle. The Botanical Review 53: 442-490
148. Sledge W. 1973. Generic and family boundaries in the Aspidiaceae and Athyriaceae. Botanical Journal of the Linnean Society 67: 203-210
149. Slosson M. 1906. How ferns grow. H. Holt & Co
150. Smith AR. 1976. Diplazium delitescens and the Neotropical species of Asplenium sect. Hymenasplenium. American Fern Journal 66: 116-120
151. Smith AR. 1990. Thelypteridaceae. In: Kramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer-Verlag
152. Smith AR. 1995. Non-molecular phylogenetic hypotheses for ferns. American Fern Journal 85: 104-122
153. Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG. 2006. A classification for extant ferns. Taxon 55: 705-731
154. Smith JE. 1793. Tentamen botanicum de Filicum generibus dorsiferarum. Mémoires de ĺAcadémie Royale des Sciences de Turin 5: 401-422
155. Stevenson D, Loconte H. 1996. Ordinal and familial relationships of pteridophyte genera. In: Camus JM, Gibby M, Johns RJ, eds. Pteridology in perspective. Kew: Royal Botanic Gardens
156. Stokey AG, Atkinson LR. 1952. The gametophyte of Blechnum spicant (L) Wither. and B. buchtienii Rosenst. Phytomorphology 2: 9-15
157. Stolze RG, Pacheco L, Øllgaard B. 1994. Polypodiaceae- Dryopteridoidea-Physematieae. In: Harling Andersson L, eds. Flora of Ecuador. Göteborg and Stockholm: University of Göteborg and Swedish Museum of Natural History
158. Stone IG. 1961. The gametophytes on the Victorian Blechnaceae. I. Blechnum nudum (Labill) Luerss. Australian Journal of Botany 9: 20-36
159. Sundue MA. 2010a.A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae). Brittonia 62: 357-399
160. Sundue MA. 2010b. A morphological cladistic analysis of Terpsichore (Polypodiaceae). Systematic Botany 35: 716-729
161. Sundue MA. 2011. Helechos y Lycophytas. In: Nee MH, ed. Flora de la Región del Parque Nacional Amboró. Santa Cruz de la Sierra, Bolivia: Editorial FAN
162. Sundue MA, Islam MB, Ranker TA. 2010. Systematics of grammitid ferns (Polypodiaceae): Using morphology and plastid sequence data to resolve the circumscriptions of Melpomene and the polyphyletic genera Lellingeria and Terpsichore. Systematic Botany 35: 701-715
163. Sundue MA, Vasco A, Moran RC. 2011. Cryptochlorophyllous spores in ferns: Nongreen spores that contain chlorophyll. International Journal of Plant Sciences 172: 1110-1119
164. Tryon AF, Lugardon. 1991. Spores of the Pteridophyta: Surface, wall structure, and diversity based on electron microscope studies. New York: Springer
165. Tryon RM. 1952. A sketch of the history of fern classification. Annals of the Missouri Botanical Garden 39: 255-262
166. Tryon RM, Tryon AF. 1982. Ferns and allied plants with special reference to tropical America. New York: Springer
167. Tryon RM, Tryon AF, Kramer KU. 1990. Pteridaceae. In: Kramer KU, Green PS, eds. The families and genera of vascular plants. Berlin: Springer-Verlag
168. Tzeng Y-H. 2002. Phylogenetic relationships of athyrioid ferns inferred from chloroplast DNA sequences. Kaohsiung, Taiwan: National Sun Yat-sen University
169. von Aderkas P. 1983. Studies of gametophytes of Matteuccia struthiopteris (ostrich fern) in nature and in culture. Canadian Journal of Botany 61: 3267-3270
170. von Aderkas P, Green P. 1986. Leaf development of the ostrich fern Matteuccia struthiopteris (L) Todaro. Botanical Journal of the Linnean Society 93: 307-321
171. Wagner WH Jr. 1952. The fern genus Diellia: Its structure, affinities and taxonomy. Berkeley and Los Angeles: University of California Press
172. Wagner WH Jr, Johnson DM. 1983. Trophopod, a commonly overlooked storage structure of potential systematic value in ferns. Taxon 32: 268-269
173. Wang M-L, Chen ZD, Zhang X-C, Lu S-G, Zhao G-F. 2003. Phylogeny of the Athyriaceae: Evidence from chloroplast trnL-F region sequences. Acta Phytotaxonomica Sinica 41: 416-426
174. Wang M-L, Hsieh Y-T, Zhao G-F. 2004. A revised subdivision of the Athyriaceae. Acta Phytotaxonomica Sinica 42: 524-527
175. Whitfield JB, Lockhart PJ. 2007. Deciphering ancient rapid radiations. Trends in Ecology & Evolution 22: 258-265
176. Wikström N, Pryer KM. 2005. Incongruence between primary sequence data and the distribution of a mitochondrial atp1 group II intron among ferns and horsetails. Molecular Phylogenetics and Evolution 36: 484-493
177. Wilson KA. 1959. Sporangia of the fern genera allied with Polypodium and Vittaria. Contributions from the Gray Herbarium 185: 97-127
178. Wolf PG, Soltis PS, Soltis DE. 1994. Phylogenetic relationships of dennstaedtioid ferns: Evidence from rbcL sequences. Molecular Phylogenetics and Evolution 3: 383-392