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Malvaceae Sl Hoehnea 31 (2): 189-214, 6 tab., 120 fig., 2004 Palinotaxonomia das tribos Byttnerieae DC., Hermannieae DC. e Helictereae DC. (Malvaceae s.l.) da flora da Bahia, Brasil 2 Marileide Dias SabaI, Francisco de Assis Ribeiro dos Santos 1,3 e Gerleni Lopes Esteves Recebido: 03.11.2003; aceito: 22.06.2004 ABSTRACT - (Palynotaxonomy ofthe tribes Byttnerieae DC., Hermannieae DC., and Helictereae DC. (Malvaceae s./.) from Bahia, Brazil). Palinotaxonomicaly, 27 species ofthe tribe Byttnerieae, Hermannieae, and Helictereae (Malvaceae s./.) from Bahia State were studied with aim ofto increase pollen morphology knowledge and to contribute with a taxonomic delimitation of the taxa. The polliniferous material were acetolysed, analyzed and photographed under light and scanning electron microscopic. Four pollen types are recognized based in exine ornamentation and apertural type: 1- (2)3(4)-porate, aspidate, planaperturate pollen grains with (micro)reticulate exine (Ayenia L., Byttneria L., and Rayleya Cristobal); II - (2)3-colporate pollen grains with microreticulate exine (Guazuma ulmifolia Lam.); III - 3 to 8-colporate pollen grains with suprareticulate or spinulose exine (Melochia L. and Waltheria L.); IV - (2)3(4)-porate, no aspidate, angulaperturate pollen grains with verrucate or spinulose exine (Helicteres L.). The results demonstrated the taxonomic importance of pollen morphology studies corroborating with the subdivision of the taxa subordinates in this tribes. Key words: pollen grains, taxonomic, Malvaceae, Bahia RESUMO - (Palinotaxonomia das tribos Byttnerieae DC., Hermannieae DC. e Helictereae DC. (Malvaceae s.I.) da flora da Bahia, Brasil). Neste trabalho e apresentado 0 estudo palinotaxonomico de 27 especies das tribos Byttnerieae, Hermannieae e Helictereae (Malvaceae s. I.) que ocorrem no estado da Bahia, com 0 objetivo de ampliar 0 conhecimento da morfologia polinica e contribuir para a melhor delimitayao dos taxons ai incluidos. Os graos de polen foram acetolisados, analisados sob microscopias optica e de varredura, descritos, fotomicrografados e eletron-micrografados. Considerando a ornamentayao da exina e 0 tipo apertural, foram reconhecidos quatro tipos polinicos: I- graos de polen (2)3(4)-porados, aspidados, planaperturados, exina (micror)reticulada (Ayenia L., Byttneria L. e Rayleya Cristobal); II - graos de polen (2)3-colporados, exina microrreticulada (Guazuma ulmifolia Lam.); III - graos de polen 3 a 8 colporados, exina suprarreticulada ou espiculada (Melochia L. e Waltheria L.); IV - graos de polen (2)3(4)-porados, nao aspidados, angulaperturados, exina verrucada ou espiculada (Helicteres L.). Pelos resultados obtidos, pode-se afirmar que a morfologia polinica tem importancia taxonomica e corrobora a subdivisao dos taxons subordinados a estas tribos. Palavras-chave: graos de polen, taxonomia, Malvaceae, Bahia Introdu~ao Manchester 1997, Bayer et al. 1999, Judd et al. 1999) nao fomecem evidencias que corroboram a subdivisao As familias Malvaceae, Bombacaceae, da ordem Malvales em quatro familias (Tiliaceae, Sterculiaceae e Tiliaceae foram situadas nos sistemas Sterculiaceae, Bombacaceae e Malvaceae), mostran­ de classificayao tradicionais (Schumann 1886, do a existencia de uma unica familia monofiletica, Dahlgren 1980, Takhtajan 1980, Cronquist 1981) na Malvaceae s.I., caracterizada principalmente pela ordem Malvales. No entanto, os limites taxonomicos presenya de um tecido nectarifero constituido de entre essas familias, bem como 0 posicionamento de tricomas glandulares, localizados na base do calice certos generos nelas incluidos, foram sempre confusos ou, menos freqUentemente, nas petalas ou no e inconsistentes, sendo questionados por varios autores androginOforo, e pelo c<ilice com preflorayaQ valvar. (Endlin 1935, Erdtman 1952). Tal circunscriyao foi adotada no presente trabalho. Estudos recentes, baseados em dados Nos sistemas de classificayao tradicionais, todos os moleculares, morfol6gicos e biogeograficos (Judd & taxons estudados aqui (27 especies, distribuidas nos 1. Universidade Estadual de Feira de Santana, Programa de P6s-Graduar;ao em Botanica, Campus Universitario, 44031-460 Feira de Santana, BA, Brasil. 2. Instituto de Botanica, Caixa Postal 4005, 01061-970 Sao Paulo, SP, Brasil. 3. Autor para correspondencia: [email protected] 190 Hoehnea 31 (2), 2004 generos Ayenia L., Byttneria L., Guazuma Plum., (1990, 1991), Roubik & Moreno (1991) e Saba & Helicteres L., Melochia L. Rayleya Cristobal e Santos (2000). Ressalta-se que as descriyoes Waltheria L.) estao situados na familia Sterculiaceae palinologicas apresentadas nesses trabalhos foram Vent. elaboradas predominantemente com base em As tribos Byttnerieae DC. e Hermannieae DC. observayoes feitas sob MO. possuem distribuiyao pantropical. Byttnerieae As especies das tribos Byttnerieae, Hermannieae caracteriza-se pelas flores com petalas unguiculadas e Helictereae foram enquadradas em tipos polinicos e cuculadas e pelo androceu formado por 5-15 estames caracteristicos, diferenciados, principalmente, pelo altemos com estaminodios (Freytag 1951, Cristobal padrao de omamentayao da exina e pelo tipo apertural, 1976), enquanto Hermannieae caracteriza-se pelas corroborando a condiyao euripolinica desses taxons flores com petalas planas e androceu formado por (Cristobal 1968, Sharma 1970). cinco estames e estaminodios reduzidos ou ausentes. No Brasil, Melhem et al. (1976) foram pioneiros, A tribo Helictereae DC. e neotropical e distingue-se no estudo dos graos de polen de especies brasileiras pela presenya de estaminodios e usualmente de urn de Helicteres, Melochia e Waltheria, elaborando uma androginOforo e pela morfologia do fruto em geral chave polinica baseada apenas em caracteres espiralado. qualitativos (tipo apertural e omamentayao da exina) A taxonomia dos generos subordinados a essas para a separayao das esp~cies. Miranda & Andrade tribos foi estudada por alguns autores, ressaltando-se (1989) tambem apresentaram uma chave polinica para as revisoes de Ayenia, Byttneria e Helicteres especies de Guazuma, Helicteres e Waltheria elaboradas por Cristobal (1960, 1976, 2001). Para incluindo, alem de caracteres qualitativos, 0 tamanho Guazuma, destaca-se 0 trabalho de Freytag (1951) e dos graos de polen. sobre Melochia e Waltheria os trabalhos de Goldberg Para a Bahia, 0 unico estudo morfopolinico sobre (1967) e Schumann (1895), respectivamente. Em representantes dessas tribos e 0 de Saba & Santos relayao as especies brasileiras, as contribuiyoes mais (2000), que analisaram seis especies da flora do Pico importantes constituem, principalmente, estudos de das Almas, municipio de Rio de Contas. Esses autores floras locais, como as de Cristobal (1983) para Santa descreveram, detalhadamente, os caracteres da exina Catarina, Costa (1981) para a regiao Sudeste do e apresentaram uma chave polinica com base em Brasil, Esteves (1986) para a Serra do Cipo e a de caracteres polinicos qualitativos e quantitativos para Cristobal et al. (1995) para 0 Pico das Almas. as especies estudadas. As contribuiyoes palinologicas mais significativas o uso da morfologia polinica na diferenciayao para os representantes das tribos aqui estudadas sao taxonomica no nivel infragenerico e de especial a de Kohler (1976) para Waltheria e a de Pire & importancia nos grupos euripolinicos (Melhem 1978). Cristobal (200 I) para Helicteres. Em seu trabalho Erdtman (1952) e Melhem (1978) referem varios sobre 0 dimorfismo polinico em Waltheria, Kohler generos aqui estudados como euripolinicos, cujos (1976) descreveu detalhadamente, sob microscopias graos de polen apresentam variayao na omamentayao, optica (MO) e eletronica de varredura (MEV), os numero, tipo e disposiyao das aberturas; isso ratifica graos de polen das flores brevistilas e longistilas e com a necessidade da ampliayao do conhecimento base em caracteres qualitativos e quantitativos palinologico dos representantes desses generos. estabeleceu dois tipos polinicos associados a o dimorfismo polinico em Melochia e Waltheria heterostilia desse genero. Pire & Cristobal (200 I), alem foi observado por Kohler (1973, 1976), Dorr & Barnett de agruparem as especies de Helicteres em nove tipos (1989) e Saunders (1993), estando, segundo esses polinicos, com base na escultura da exina (MO e autores, associado a heterostilia caracteristica de suas MEV), analisaram as afinidades taxonomicas especies. Segundo Saxena (1993), isso pode existentes entre esse genera e os outros da tribo representar urn dos fatores morfologicos distintivo dos Helictereae. sistemas de incompatibilidade de plantas heterostilicas. Outros trabalhos importantes sobre a morfologia o presente trabalho teve como objetivo analisar polinica de generos das tribos estudadas sao: Erdtman a morfologia polinica de especies das tribos (1952), Cristobal (1968), Chaudhuri (1969), Sharma Byttnerieae, Hermannieae e Helictereae (Malvaceae (1970), Kohler (1971,1973), Melhem et al. (1976), s.l.) ocorrentes na flora da Bahia, a fim de ampliar os Miranda & Andrade (1989), Palacios-Chavez et al. conhecimentos sobre a morfologia polinica e fomecer M.D. Saba, F.A.R. Santos & G.L. Esteves: Palinotaxonornia de Malvaceae s.l. da Bahia 191 dados que auxiliem na delimitayao taxonomica dessas A terminologia palinologica utilizada foi a de Punt tribos e dos taxons nelas incluidos. et al. (1994). A nomenclatura de Barth & Melhem (1988) foi tambem utilizada, mas apenas como apoio Material e metodos para transcriyao de alguns termos para 0 portugues. Com base em levantamento de herbario e de Material examinado:
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