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303 T07 21 07 2015.Pdf lIoehnea 30(3): 243-283, 2 tab., II fig., 2003 Grammitidaceae (Pteridophyta) no Brasil com enfase nos generos Ceradenia, Cochlidium e Gramlnitis 2 Paulo Henrique Labiak 1,3 e Jefferson Prad0 Reccbido: 18.06.2003: aceito: 28.10.2003 ABSTRACT - (Grammitidaceae (Pteridophyta) from Brazil with emphasis on the genera Ceradenia, Cochlidiul11 and Cral11l11itis). In this work is presented a taxonomic revision for the species ofthe genera Ceradenia [CO albidula (Baker) L.E. Bishop, C. capillaris (Desv.) L.E. Bishop, C. gla~iovii (Baker) Labiak, c.jungerl11anioides (Klotzsch) L.E. Bishop, C. pruinosa (Maxon) L.E. Bishop, C. spixiana (Mart. ex Mett.) L.E. Bishop, C. warl11ingii (c. Chr.) Labiak], Cochlidiul11 [C .jurcatul11 (Hook. & Grev.) C. Chr., C. linearifoliul11 (Desv.) Maxon ex C. Chr., C. pUl11ilul11 C Chr., C. punctatul11 (Raddi) L.E. Bishop, C. serrulatul11 (Sw.) L.E. Bishop, C. tepuiense (A.C. Smith) L.E. Bishop], and Cral11l11itis [Gjlul11inensis Fee, G leplopoda (CH. Wrigth) Copel.] which occur in Brazil, with considerations about classification, morphology, and geographical distribution ofthe family, as well as identification keys for all the genera in Brazil. For the taxa here treated, are also provided identification keys, descriptions, comments, illustrations, and geographic distribution. Key words: flora, pteridophytes, revision, taxonomy RESUMO - (Grammitidaceae (Pteridophyta) no Brasil com enfase nos generos Ceradenia, Cochlidium e Cral11milis). Este trabalho apresenta uma revisao taxonomica das especies dos generos Ceradenia [CO albidula (Baker) L.E. Bishop, C. capillaris (Desv.) L.E. Bishop, C. gla~iovii (Baker) Labiak, C. jungerl11anioides (Klotzsch) L.E. Bishop, C. pruinosa (Maxon) L.E. Bishop, C. spixiana (Mart. ex Mett.) L.E. Bishop, C. warl11ingii (C. Chr.) Labiak], Cochlidium [C .jurcatum (Hook. & Grev.) C. Chr., C. Iinearifoliul11 (Desv.) Maxon ex C. Chr., C. pUl11ilum C Chr., C. punctatul11 (Raddi) L.E. Bishop, C. serrulatul11 (Sw.) L.E. Bishop, C. tepuiense (A.C. Smith) L.E. Bishop] e Cral11l11itis [Gjlul11inensis Fee, G leptopoda (C.H. Wrigth) Copel.] que ocorrem no Brasil, alem de considerac;5es sobre 0 hist6rico de classificac;ao, aspectos sobre morfologia e distribuic;ao geografica da familia, bem como chaves de identificac;ao para todos os generos de Grammitidaceae no Brasil. Para as especies aqui consideradas, tambem sao fornecidas chaves de identificac;ao, descric;5es, comentarios, ilustrac;5es e distribuic;ao geografica. Palavras-chave: flora, pterid6fitas, revisao, taxonomia Introdu~ao Grammitidae. Posteriormente, Hooker (1844-1864) as incluiu na Tribo Grammitideae, como parte da A familia Grammitidaceae compreende cerca de Subordem Polypodiaceae. Autores subseqUentes, no 500 especies, distribuidas em aproximadamente 15 entanto, optaram pOl' tratar esses taxons em um sentido generos, presentes nos tr6picos e subtr6picos de todo amplo, mantendo-os em Polypodiaceae s.l. (Mettenius o globo. Na regiao Neotropical esta representada pOl' 1856, Baker 1870, Engler & Prantl 1898-1902). nove generos e ca. de 200 especies, contemplando Christensen (1938), ao trabalhar com a familia quase que a metade das especies existentes (Moran Polypodiaceae s./., chamoLl a atencyao sobre a et al. 1995). necessidade de estudos mais detalhados sobre os as representantes da familia Grammitidaceae diferentes grupos que a constituiam, com a finalidade eram reconhecidos classicamente como um grupo de se criar LIm sistema mais natural de organizacyao. pertencente a Polypodiaceae s.!., agrLlpados em Finalmente, em seu trabalho "On natural classification subfamilias ou tribos. Presl (1836) foi 0 primeiro autor ofthe family Polypodiaceae", Ching (1940) propos 0 a reunir os representantes de Grammitidaceae reconhecimento de Grammitaceae, com duas tribos, estabelecendo a Tribo Grammitaceae, com LIma Secyao reunindo aproximadamente 400 especies, Grammitieae I. Universidade Federal do Parana. Setor de Cieneias Biol6gieas. Departamento de Botiiniea, Caixa Postal 1644,80011-990 Curitiba. PRo Brasil. 2. Instituto de Botiiniea. Sec,:ao de Briologia e Pteridologia. Caixa Postal 4005, 0 I061-970 Sao Paulo. SP. Brasil. 3. Autor para eorrespondeneia: [email protected] 244 Hoehnea 30(3), 2003 (Grammitis SW., Calymmodon C. Presl, Acrosorus caule e curtamente ascendente ou ereto, com as Copel., Prosaptia C. Presl e Holcosorus T. Moore) frondes nao-afiiculadas com 0 mesmo, pedicelo do e Cochlideae (Cochlidiul11 Kaulf., Sc!eroglossum esporangio com apenas uma fileira de celulas, esporos A1derw., Nematopteris Alderw. e Oreogrammitis triletes clorofi1ados e gamet6fitos filamentosos, Copel.). ramificados. Os esporos de Grammitidaceae apresen­ o nome da familia Grammitaceae, erroneamente tam exce<;:oes para algumas especies, podendo ser grafado par Ching (1940), foi corrigido par Holttum monoletes; porem, sempre c1orofilados (Wagner 1985). (1947), uma vez que a con'eta ortografia deve ser Estudos filogeneticos envolvendo as duas familias formada a pafiir do genitivo singular do nome generico sao ainda bastante escassos. Dados preliminares ("Grammitidis"), passando entao para Grammitidaceae apresentados par Pryer et al. (1995), obtidos a pafiir (Artigo 18.1 do C6digo Internacional de Nomenclatura da analise filogenetica baseada em caracteres BoHinica, Greuter et al. 2000). Esta grafia vem sendo morfol6gicos e molecu1ares (rbcL) para diversos seguida pelos autores subseqUentes, exceto pOI' Pichi­ grupos de pterid6fitas, mostram-se distintos conforme Sermolli (1970) que, embora tenha comentado sobre a analise, combinada ou nao, dos caracteres a mudan<;:a proposta por Holttum (1947), ainda utilizou morfol6gicos e moleculares. Na analise de maxima a grafia Grammitaceae. parcim6nia utilizando-se apenas os caracteres o conceito de Ching (1940), no entanto, nao foi morfol6gicos, Grammitidaceae (representada pOI' de todo aceito pelos estudiosos do tema, como pOI' Micropolypodium Hayata) mostra-se mais exemplo Copeland (1947), que em sua obra "Genera estreitamente relacionada aos generos Vittaria Sm. Filicum" manteve 0 genero Grammitis na familia (Vittariaceae) e Loxogramme (Blume) C. Presl Polypodiaceae. A proposta de c1assifica<;:ao de Ching (Polypodiaceae), formando um grupo parafil6tico com (1940) teve maior impulso apenas quando Pichi­ Polypodiaceae (representada pOI' Polypodium L. s.s.). Sermolli (1958) dividiu Polypodiaceaes. f. em diversas De maneira oposta, na analise de maxima parcim6nia familias, dentre as quais figuravam novamente as baseada apenas nos caracteres moleculares, 0 genera Grammitidaceae (citada como Grammitaceae). Micropolypodium aparece no mesmo clado de Recentemente, as Grammitidaceae vem Polypodium, formando assim um grupo monofiletico. recebendo maior aten<;:ao, evidenciada principalmente Essa rela<;:ao e mantida pela analise combinada dos nos trabalhos realizados pOI' Bishop (1974, 1977, 1978, caracteres morfol6gicos e moleculares, apenas com 1988, 1989a, b), com as especies do novo mundo, uma diminui<;:ao no indice de "bootstrap". Parris (1983, 1984, 1986, 1990, 1995, 1997a, 1998) e Deve-se ressaltar que, embora as duas familias Parris et af. (1992), com as do velho mundo. Nesses provavelmente sejam de origem monofiletica, estudos trabalhos, Bishop iniciou um estudo mais minucioso mais criteriosos, com a inclusao de outros taxons e de sobre os diferentes grupos encontrados dentro das novos caracteres morfol6gicos e moleculares, sao Grammitidaceae, reconhecendo varios generos. necessarios para 0 reconhecimento das provaveis Infelizmente Bishop veio a falecer antes de completar rela<;:oes filogeneticas entre ambas, e 0 reconhecimento seus estudos, os quais foram em paJie concluidos pOI' das mesmas como taxons distintos ou nao. Smith e Moran, alguns a partir dos seus pr6prios Dos 15 generos atualmente reconhecidos em manuscritos (Smith et al. 1991, Bishop & Smith 1992, Grammitidaceae, nove ocorrem na regiao neotropical, Smith 1992, Smith & Moran 1992, 1993a, b,). sendo eles: Ceradenia L.E. Bishop, Cochlidium As rela<;:oes da familia Grammitidaceae com as Kaulf., Enterosora Baker, Grammitis, Lellingeria Polypodiaceae s. s. tem sido ainda amplamente discu­ A.R. Sm. & R. C. Moran, Melpomene A.R. Sm. & tidas, principalmente porque em habito, divisao da R.C. Moran, Micropolypodium, Terpsichore A.R. lamina e ausencia de indusio, algumas especies de Sm. e Zygophlebia L.E. Bishop. Desses, apenas Grammitidaceae lembram especies de Polypodiul11 Enterosora nao foi ainda registrado para 0 Brasil, em miniatura. As Po1ypodiaceae, porem, podem ser apesar de apresentar distribui<;:ao geognlfica bastante diferenciadas pelo caule claramente decumbente, com ampla, ocorrendo nos paises da America Central e as frondes afiiculadas e inseridas dorsalmente sobre America do Sui ocidental, em regioes muito pr6ximas o caule em duas fileiras, pedicelo do esporangio com as fronteiras do Brasil. duas fileiras de celulas, esporos monoletes aclorofilados Estudos envolvendo as Grammitidaceae s.s. do e gamet6fitos cordiformes. Em Grammitidaceae 0 Brasil ainda sao escassos, embora algumas das P.I-!. Labiak, J. Prado: Os gencros Ceradel1ia, CochlidiulIl c Gralllmilis no Brasil 245 especies que ocorrem no Brasil estejam incluidas em BR, FLOR, GH, GUA, HB, HBR, HRCB, INPA, IPA, trabalhos realizados para outras regioes das Americas, K, MBM, NY, OURP, PACA, PEUFR, R, RB, SJRP, como pOl' exemplo Copeland (1952a, b, 1956) para as SP, SPF, UC, UEC, UFJF, UFP, UPCB e US. Americas em geral; de la Sota (1966b) de la Sota Encontram-se citados os materiais brasileiros et al.
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