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Significance of Gametophyte Form in Tropical, Epiphytic Ferns Cynthia Lynn Dassler Iowa State University

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Significance of Gametophyte Form in Tropical, Epiphytic Ferns Cynthia Lynn Dassler Iowa State University Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1995 Significance of gametophyte form in tropical, epiphytic ferns Cynthia Lynn Dassler Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons, and the Ecology and Evolutionary Biology Commons Recommended Citation Dassler, Cynthia Lynn, "Significance of gametophyte form in tropical, epiphytic ferns " (1995). Retrospective Theses and Dissertations. 10774. https://lib.dr.iastate.edu/rtd/10774 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directfy from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from ai^ ^pe of coiiq}uter printer. The qnality of this reproduction is dqoendait upon the quality of the copy snbmitted. Broken or indistinct print, colored or poor quality illustrations and photogrs^hs, print bleedthrough, substandard margins, and inqvoper alignment can adversety affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note win indicate the deletion. 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Ml 48106-1346 USA 313.'761-4700 800/521-0600 Significance of gametophyte form in tropical, epiphytic fems by Cynthia Lynn Dassler A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Department: Botany Major: Botany (Systematics and Evolution) Approved: Signature was redacted for privacy. In Charge of Major Work Signature was redacted for privacy. For the Major Departmeut Signature was redacted for privacy. For the Graduate College Iowa State University Ames, Iowa 1995 UMI Number: 9540888 OHI Microform 9540888 Copyright 1995, by UMI Company. All rights reserved. 1!his microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, HI 48103 ii For my mother, Jacqueline Jo Dassler, for her inspiration that has continually been my guide. and For Bear, whose friendship, love and enjoyment of life gave and continue to give me courage. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS iv GENERAL INTRODUCTION I 1. GAMETOPHYTE FORM, LONGEVITY AND HABITAT 8 2. SIGNIFICANCE OF FORM IN FERN GAMETOPHYTES: CLONAL, GEMMIFEROUS 55 GAMETOPHYTES OF CALLISTOPTERIS BAUERANIA (HYMENOPHYLLACEAE) 3. GERMINATION AND SURVIVAL OF EPIPHYTIC AND TERRESTRIAL FERN 99 GAMETOPHYTES IN BRYOPHYTE MATS 4. SIGNIFICANCE OF GREEN SPORES IN EPIPHYTIC FERNS 116 5. SIGNIFICANCE OF GAMETOPHYTE FORM IN LONG DISTANCE COLONIZATION 132 BY TROPICAL, EPIPHYTIC FERNS 6. COMBINING INDEPENDENT GAMETOPHYTE POPULATIONS 169 GENERAL CONCLUSIONS 176 iv ACKNOWLEDGEMENTS I would like to thank Dr. Don Farrar for stimulating ideas and discussions, sharing his love of nature and his example as a teacher. Thanks also go to my committee. Dr. Lynn Clark, Dr. Nels Lersten, Dr. Robert Wallace, Dr. George Brown, and Dr. Edwin Powell, for their ideas and support. My mother, Jacqueline Dassler, and sister, Deborah Dassler, gave me love, encouragement and support throughout; I could not have done this without them. Very special thanks go to Chris Richardson for everything. Various portions of this work were supported by the Iowa State University Pohl Fund, an Iowa State University Professional Advancement Grant, an American Society of Plant Taxonomists grant for student research and a Sigma Xi Grant-In-Aid of Research. 1 GENERAL INTRODUCTION Introduction to Hypotheses A full understanding of evolutionary relationships and directions of evolutionary change within a plant group requires knowledge of the morphological and ecological adaptations of both the gametophyte and sporophyte generations. In ferns, where the sporophyte and gametophyte live independently, the morphology and ecology of the sporophyte has been extensively studied. Most fern species have been described and classified according to sporophyte characters and this is reflected in the characters used in construction of fern phylogenies. Studies of the gametophyte generation, however, are relatively few and generally are not used in evolutionary hypotheses. The typical fern gametophyte described in textbooks is small, thallose, heart-shaped and short-lived, usually dying in a few months' time whether or not a sporophyte is produced This description, however, does not convey the true complexity and variation of fern gametophytes. Gametophytes possess different morphologies which suggest different environmental adaptations. Some gametophytes grow underground and are not green and photosynthetic. Among green photosynthetic fern gametophytes, some are heart-shaped and some are ribbon-like, strap-shaped or filamentous. Nonheart-shaped, green gametophytes are long-lived and clonal. In addition, some of these gametophytes produce gemmae at the tips of the thallus or filament. Thus, instead of one category of green fern gametophytes, as presented in most texts, there are three types: 1. Short-lived gametophytes, which occur in most terrestrial ferns. 2. Persistent, clone-forming gametophytes lacking gemmae, which are found in Polypodiaceae and Elaphoglossaceae. 3. Persistent, clone-forming gametophytes possessing gemmae, which are found in Hymenophyllaceae, Grammitidaceae and Vittariaceae. Persistent, clone-forming gametophytes (i.e., in the five families Polypodiaceae, Elaphoglossaceae, Hymenophyllaceae, Grammitidaceae and Vittariaceae) are common in tropical, epiphytic habitats. Literature on gametophytes of these families has consisted of a relatively few papers which extend little beyond the development 2 and morphological description of the gametophyte (Nayar and Kaur 1971). Research on the evolution of morphologies and ecological adaptations of tropical epiphytic gametophytes is very limited. Thus, several of the techniques and experiments in this research are completely new. Some worked, and some did not. Questions addressed in this research include; 1. Why do gametophytes of epiphytic species possess persistent, clone-forming growth? 2. What is it about a tropical epiphytic habitat that has driven evolutionarily unrelated families to evolve similar gametophyte growth forms? 3. What is the role of gemmae? Questions such as these led me to Hawaii to observe gametophytes in their natural habitats. In Hawaii, these gametophytes were observed growing, with or without associated sporophytes, as mats, sometimes extensive ones, with intermixed bryophytes, or often interwoven and tangled within the bryophyte mat in epiphytic habitats. These observations led to the first of two hypotheses: Hypothesis One: Persistent gametophyte growth forms in ferns evolved to compete with bryophytes in the epiphytic habitat by increasing the probability of the gametophytes finding a favorable space and/or time for the production of sporophytes. Persistent gametophyte growth increases the opportunity for a gametophyte to find a microsite favorable for growth and reproduction, as well as to survive longer and wait for the arrival of another gametophyte so that outbreeding can occur. The question remains as to whether persistent growth is enough to bring two different clones together to facilitate outbreeding, or are epiphytic ferns mostly inbreeders? There are two ways to examine the question of whether epiphytic ferns are inbreeders or outbreeders. These are examination of the genetic structure of sporophyte populations and examination of biological adaptations of the gametophyte that influence breeding systems. The genetic structure of sporophyte populations, as determined by enzyme electrophoresis, indicates that most ferns, including epiphytes, are outbreeders (Soltis and Soltis 1989,1992; Ranker 1992; Haufler et al. 1995). Biological adaptations, specifically the presence of an antheridiogen system, also indicate that epiphytes are outbreeders. An antheridiogen system promotes outcrossing by using chemical means to insure that the sexes are separated onto two different gametophytes. All homosporous 3 fem gametophytes have the potential to become bisexual and thus inbreed. Self-fertilization of a bisexual gametophyte results in totally homozygous sporophytes, and the possibility of inbreeding depression and expression of deleterious alleles. In fem species with antheridiogen systems, the faster-growing
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