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Taxonomy, Distribution, and Seasonal Occurrence of the Saprolegniaceae in Aquatic Sites in Northwest Iowa Maren A Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1980 Taxonomy, distribution, and seasonal occurrence of the Saprolegniaceae in aquatic sites in northwest Iowa Maren A. Klich Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons Recommended Citation Klich, Maren A., "Taxonomy, distribution, and seasonal occurrence of the Saprolegniaceae in aquatic sites in northwest Iowa " (1980). Retrospective Theses and Dissertations. 7339. https://lib.dr.iastate.edu/rtd/7339 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 was produced from a copy of a document sent to us for microfilming. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the material submitted. The following explanation of techniques is provided to help you understand markings or notations which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure you of complete continuity. 2. When an image on the film is obliterated with a round black mark it is an indication that the film inspector noticed either blurred copy because of movement during exposure, or duplicate copy. Unless we meant to delete copyrighted materials that should not have been filmed, you will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photo­ graphed the photographer has followed a defînite method in "sectioning" the material. It is customary to begin filming at the upper left hand comer of a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again—beginning below the first row and continuing on until complete. 4. For any illustrations that cannot be reproduced satisfactorily by xerography, photographic prints can be purchased at additional cost and tipped into your xerographic copy. Requests can be made to our Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In all cases we have filmed the best available copy. Universi^ MicrOTlms International 300 N, ZEEB ROAD, ANN ARBOR, Ml 48106 18 BEDFORD ROW, LONDON WC1R 4EJ, ENGLAND 8012972 KUCH, MAREN A. TAXONOMY. DISTRIBUTION, AND SEASONAL OCCURRENCE OF THE SAPROLEGNIACEAE IN AQUATIC SITES IN NORTHWEST IOWA Iowa State University PH.D. 1980 University Microfilms IntGrnStiOnSi 300 N. Zeeb Road. Ann Arbor, MI 48106 18 Bedford Row, London WCIR 4EJ, England Copyright 1980 by Klich, Maren A. All Rights Reserved Taxonomy, distribution, and seasonal occurrence of the Saprolegniaceae in aquatic sites in northwest Iowa by Maren A. Klich A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Department: Botany Major: Botany (Mycology) Approved: Signature was redacted for privacy. In Charge of Major Work, Signature was redacted for privacy. For the Major Department Signature was redacted for privacy. For the Gradu College Iowa State University Ames, Iowa 1980 Copyright (£) Maren A. Klich, 1980. All rights reserved. ii TABLE OF CONTENTS Page GENERAL INTRODUCTION 1 PART I 3 LITERATURE REVIEW 4 Morphology 4 Collection 4 Culture 7 Ecology 10 MATERIALS AND METHODS 15 Preliminary Laboratory Studies 15 Field Sites and Sampling Techniques 17 Laboratory Culture 23 RESULTS AND DISCUSSION 24 Species Isolation 24 Effects of Cultural Regimes 26 Comparison of Field Sites 30 Seasonal Periodicity and Water Temperatures 36 SUMMARY 63 PART II 65 INTRODUCTION 66 COMPUTERIZED SYNOPTIC KEY TO SPECIES OF ACHLYA AND SAPROLEGNIA 70 LITERATURE CITED 83 ACKNOWLEDGMENTS 92 APPENDIX A. COLLECTION DATES AND WATER TEMPERATURES 93 APPENDIX B. PRECIPITATION AND TEMPERATURE DATA FOR NORTHWEST IOWA, 1976-1978 96 APPENDIX C. SIMILARITY INDICES 100 APPENDIX D. COMPUTER PROGRAM FOR A HYPOTHETICAL SYNOPTIC KEY 103 1 GENERAL INTRODUCTION The Oomycete family Saprolegniaceae is a group of predominantly aquatic fungi characterized by biflagellate zoospores. Although these fungi are common in freshwater habitats, their role in aquatic ecosys­ tems is virtually unknown. This may be due largely to isolation and identification difficulties. The purposes of this work were 1) to study the occurrence and distribution of the aquatic species of the Sapro­ legniaceae in northwest Iowa, 2) to refine isolation and culture methods for these fungi, 3) to devise a workable alternative to existing dichot- omous identification keys for species belonging to the common aquatic genera Achlya and Saprolegnia. Some members of the Saprolegniaceae reportedly display a distinct seasonal occurrence (Coker 1923, Hughes 1962, Dick and Newby 1961, Roberts 1963, Alabi 1971a,b). The field study was designed to investi­ gate this ecological aspect of the family by repeatedly sampling several aquatic sites for two years. Differences in the fungal flora between sites were also studied. To expedite ecological work with the Saprolegniaceae, techniques for isolation and culture should be as simple as possible. The tradi­ tional isolation substrate has been boiled hemp (Cannabis sativa L.) seeds. These seeds are often not readily accessible, therefore, several more common seeds were compared with hemp seeds as isolation substrates. The identification process for species of Saprolegniaceae is tedious at best. Every isolate must be observed several times over 2 one to three weeks to study taxonomically important characters as they develop. Some isolates never produce the structures necessary for identification utilized in standard dichotomous keys, but may possess other characters that could aid in identification. The various sorting systems and random-access keys allow workers to use several characters to identify an organism, but these keys have not gained acceptance because they are unwieldy. The synoptic key included in the final section of this paper has been computerized for easy use. 3 PART I 4 LITERATURE REVIEW Morphology The Saprolegniaceae is characterized by a vegetative thallus com­ posed of coenocytic hyphae. The vegetative colony of most aquatic species is 1-5 cm in diameter on a hemp seed substrate. The vegetative mycelium, once believed to be haploid (Patterson 1927, Shanor 1937), is now generally considered to be diploid (Heath and Greenwood 1968, 1970; Howard and Moore 1970). Sexual reproduction is oogamous, with one to many oospheres per oogonium. Antheridial hyphae attach to the oogonial wall and form fertilization tubes through which a single male nucleus passes to fertilize each oosphere. Two major forms of asexual repro­ duction exist within the family. The first is by production of zoospores characterized by possessing one tinsel type and one whiplash flagellum. Some members of the family are diplanetic with pip-shaped primary zoo­ spores and reniform secondary zoospores. The second type of asexual reproduction is by formation of chlamydospores or gemmae. These gemmae may eventually develop into zoosporangia, oogonia with oospores develop­ ing parthenogenetically, or germinate directly by a germ tube. A good account of the morphological characteristics of the family may be found in Dick (1973). Collection Aquatic saprolegniaceous fungi are inconspicuous and ephemeral and, therefore, cannot easily be collected from the environment as one would collect vascular plants or mushrooms. Generally, a suitable 5 substrate is placed in the water, or in a water sample in the laboratory, from which one wishes to isolate species of the family. These sub­ strates, appropriately referred to as "baits," are colonized by fungal propagules in the water. The resultant fungal thalli can then be observed and identified. To those not familiar with the baiting techniques used for aquatic fungi, a list of isolation substrates would appear to be more appropriate as a recipe for a witch's brew than for any scientific purpose. Among the baits used are dead flies, termites, ants, ant pupae, snake skin, grass blades, mushroom grubs, egg yolk, and various kinds of fruits, twigs and seeds (Pieters 1915, Coker 1923, Harvey 1925, Cejp 1934, Wolf and Wolf 1941, Hamid 1942, Beneke 1948a, Hoffman 1949, Johnson 1956, Cooke and Bartsch 1960, Scott 1961, Dick 1966, Thakur Ji 1970, Alabi 1971a, Rooney and McKnight 1972, Zebrowska 1976, Padgett 1978a,b). Various workers have used semi-solid media, but Ho (1975) developed a medium that is reportedly selective for Saprolegnia species. Although Scott (1961) found boiled snake skin to be the best bait for Aphanomyces, most workers have found boiled hemp seed preferable for most members of the Saprolegniaceae. Zebrowska (1976) compared several isolation sub­ strates and found that saprolegniaceous fungi appeared more frequently on hemp than on flies, fruits, or twigs. Harvey (1925) noted that
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