Life History and Reproductive Biology of the Ferns of Woodman Hollow, Webster County, Iowa James Herbert Peck Iowa State University
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Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1980 Life history and reproductive biology of the ferns of Woodman Hollow, Webster County, Iowa James Herbert Peck Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons Recommended Citation Peck, James Herbert, "Life history and reproductive biology of the ferns of Woodman Hollow, Webster County, Iowa " (1980). Retrospective Theses and Dissertations. 6799. https://lib.dr.iastate.edu/rtd/6799 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]. 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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 CODV. UnK^fS^v' /VAirrrvfJImc internarioncii • e SEDFORD ROW, LONDON WCl A ENGLAND 8028628 PECK, JAMES HERBERT LIFE HISTORY AND REPRODUCTIVE BIOLOGY OF THE FERNS OF WOODMAN HOLLOW, WEBSTER COUNTY, IOWA lo-wa Slate University PH.D. 1980 university MIcrofiims 11 IICI I ISLIUI ISL 300 N. Zeeb Road, Ann Arbor, MI 481CK) iS Bedford Row, London WCIR 4EJ, England Copyright 1980 by Peck, James Herbert All Rights Reserved PLEASE NOTE: In alt cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been Identified here witn a cneck mark . 1. Glossy photographs •/ 2. Colored illustrations 3. Photographs with dark background '4. Illustrations are poor copy _______ 5. °rint shows through as there is text on both sides of page ]y^ 5. Indistinct, broken or small print on several pages throughout 7. Tightly bound copy with print lost in spine 8. Computer printout pages with indistinct print 9. Pcce(s) lacking when material received, and not available from school or author 10. Page(s) seem to be missing in numbering only as text fcl1ows 11. Poor carbon copy 12. Not original copy, several pages with blurred type 13. Anpendix pages are poor copy 14. Original copy with light type 15. Curling and wrinkled pages _____ 16. Other Jrll v'SfSTTj/ Mkzonims ritsrr^xioncil Life history and reproductive "biology of the ferns of Woodman Hollow, Webster County, Iowa Dy James Herbert Peck A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Major: Botany ATDDrovea: Members of the Committee: Signature was redacted for privacy. Signature was redacted for privacy. Signature was redacted for privacy. [n Charge of Major Work Signature was redacted for privacy. Signature was redacted for privacy. Signature was redacted for privacy. ^or :he Major DeparisiepfC Signature was redacted for privacy. Signature was redacted for privacy. For the Graduate Cbllege LVAa O ucv vc OAi-LV S^o-1-u.y Ar.es. Iowa 1 oAn n 0. T___ .J. T. 4 rvO/-N ow I-'V J. uea-iiiCO ii' era. »-/rr— u i ov-rw • J- , il iii TABLE OF CONTENTS Page IN M3M0RIAM v INTRODUCTION 1 PART I. INVESTIGATIONS OF STUDY AREA ECOLOGY 11 INTRODUCTION 1] NATURAL FEATURES 15 POST-SETTLEMENT HISTORY 3l POST-PLEISTOCENE PHYTOGEOGRAPHY 35 MICRQGLÏMA1ES 4-5 BRYOFLORISnC ANALYSIS 83 PART II - INVESTIGATIONS ON SPOROPHYîE ECOLOGY 95 INTRODUCTION 97 PROCEDURES 99 IDENTIFICATIONS I03 FLORISTIC RELATIONSHIPS 11? DISTRIBUTION WITHIN STUDY AREA I35 FOLIAR PHENOLOGY 157 PERMANENT STUDY PLOTS 173 PART III. INVESTIGATIONS ON SPORE ECOLOGY 199 INTRODUCTION 201 uncio 03 IDELFTIFICATIONS 213 PRODUCTION 239 RELEASE 247 DISPERSAL 269 GERÎ'ÎINATION 33I PART IVI II'T.'IS'RJ.GA'J.-I.'W.'RIS 'JR- UÏ'IZJ ILCJWLCGX 3^7 INTRODUCTION 349 PROCEDURES 353 IDEITTIFICATIOINS 361 OCCURRENCE 377 DEMOGRAPHICS 3S9 KGRTALITY 399 IV SEXUALITY 417 PERIODICITY 433 PART V. INVESTIGATIONS ON SPORELING ECOLOGY 441 INTRODUCTION 443 PROCEDURES 4^1-5 IDENTIFICATIONS 449 PRODUCTION 461 MORTALITY 46? MATURATION 473 DISCUSSION 483 SUMMARY 503 REFERENCES 523 ACKNOWLEDGEMENTS 543 V IN MEMGRIAM Dr. Ludwig F. Guldner, M. D. (1881-1974), a prominent physician in ïïiy home +<5-^,. Davenport ; lo"?.; received, his medical degree from the University of Iowa in 1902 and studied in Vienna for two years "before returning to Davenport to practice. While abroad, he developed a strong interest in floristic botany. That interest lay dormant while Dr. Guldner established the local medical pathology laboratory, authored several textbooks on pathology, and performed the duties of county coroner. After 52 years of service, he retired from medicine and renewed his study of flcristic botany. I met Dr. Guldner in 195? at a time •s-ihen he "K&S ooaplstlng a 20 year- stud;y of the flora of Scott and Muscatine counties. He introduced me to botanical keys and helped me leam the names of the local plants. From these trips and keying sessions, I acquired an appreciation of the plants and their habitats, as well as, a sense of tragedy of their continued extirpation. As an expression of gratitude, this dissertation is dedicated +-o Dr. Guldner. 1 INTRODUCTION Statement of the Problem Detailed observations on life history and reproductive biology have been made on few vascular plants. Most were conducted on annual plants common to managed ecosystems, rather than on perennial, herbaceous plants, such as ferns (Harper, 1977)• The study of fem life history and reproductive biology is potentially even more complex tl-ian that encountered with seed plants. The fem life cycle consists of two, independent generations (Figure 1), rather than the sporophyte dependent life cycle of seed plants. It is, therefore, not entirely surprising that life history and reproductive studies on ferns in nature are exceedingly rare, since twice as much effort might need to be expended per species. The readily observed alternation of the two generations of a fem life cycle has provided generations of botanists with a useful laboratory tool to investigate significant problems in biology (Whittier, 1971) • Several thousand laboratory investigations on gametophytes are reviewed by Atkinson (1973) > Lloyd (19? 4-), Killer (1968), Nàf _et al.