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Submitted to the Faculty a Thesis • of of Jo Ann Banks August 19]8

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Submitted to the Faculty a Thesis • of of Jo Ann Banks August 19]8 ____...IIII:~·i::s;c:.: ' THE BIOSYSTEMATICS :OF ERYTHRONIUM PROPULLANS GRAY AND SYMPATRIC POPULATIONS OF ERYTHRONIUt~ ALBIDU~1 NUTTALL (LILIACEAE) AThesis Submitted to the .. Faculty of University of Wisconsin" LaCrosse La Crosse, Wisconsin 54601 Jo Ann Banks lriPartial Fulfillment bfthe Requirements for the Degree • of Master of Science in Biology August 19]8 © Mfr­ -1<,/; 8tL'n UNIVERSITY OF WISCONSIN .. LA CROSSE La Crosse, Wisconsin 54601 COLLEGE OF ARTS, LETTERS, AND SCIENCES Candidate: Jo Ann Banks We recommend acceptance of this thesi.sto. theColl ~g~ofArts.'. L.etters, and Sciences in partial fulfillment of this candidates' requirements for the degree Master of Science in Biology. The candidate has completed her oral defense of the thesis. 'l3i.?Y Date ~ ,-31-7!- ::::::::r; '. ~ y~ ~':::? - .v"=" .JI.. zr~--'- ~.- Datj,b '61(g/&a: ~J'}~p:~ ../t)at-e This thesis is app"oved for the College of Arts, Letters, 79-02634 ii ABSTRACT A biosystematic.comparison of Erythroniumpropullans.ilndsympatric populations of f. albiQum was conducted to clarify the relationships of these •.. twospecte.s.. .iE.propu11 ans . iSiendemicto,southeaste.rn.Minnespta., while I. albidum is common and widespread throughout the eastern United States, including southeastern ~1i nnesota. Themorphol ogy,geography, .cy­ tology, ecology, and reproductive biology of each species were investigated. Populations of E. albidum and E. propullans within Rice and Goodhue Counties, Minnesota, were utilfzedinthesttidY.. A morphological analysis of six. charactersjndicateclthatt. albidum and I. propullans are morphologically distinct species, and that th,ree size classes of plants exist: f. propull ans, depictedasa.dwarfed f. ~­ bidum, an intermediate size class, and L al:!~jdum. The intermediate size class suggested that La1bidum andE:proplrl-ra:ns--hyhri-d-Fze-te--formferti le offspring. The rari ty.of the hybridfurther.suggested that sOme repro- ductive isolation between the two species exists. The pollen-ovule ratio, protandry, and the dimorphi c stamens of I. propull ansand I .. al bi dum suggest that both species are facultatively outcrossing. Breeding experiments demonStrated that I. albidum is facultatively outcrossing and able to pro~ duceseed when. crosseclwith I. propul1etDS_. Selt-ferti1i zationi~E.a 1­ bidum required a physical mechanism to transfer pollen f~oril-tlieantherto the. stigma. E. propull ans, however, produced seed only when crossed with E. albidum. Pollen .was found to be effectively transferred either between C10rle?of I. albidum ()Y' f. propullans by AndrenacarliniCkl1., a solitary oligolectic bee whose pollination behavior waswells~i~~<i~()thefl()wers of. both species . Vi sits. by this i nsectwer~ •• PY'iJTI~rilYi~oiE •.• al bi dum flowers. The l~eproductive evidence questions the validitYiof the spectfic ranking gi ven to Erythroni um propull ans as suggested'b,y its morpho logy. The re­ stricted range, the relative sterility,. and.the .• ploidyl/ev.elof.E.pro­ pul+ans suggest it isarelatively youngspeciesofrecentori gi n derived from f. albidum. The inability of I. propullanstoreproducewith and a natural low level of seed production in this species, further suggested that populations are maintained exclusively by vegetative re­ production. The lack of input of genetic variability by sexual processes indicates that I. Qropullans will not adapt to drastically changing en­ \liJ·QnlJle[);t~.~~JJnle_s~~=S1:ttj<:a1 habitat ispr~s~Y'ye<i, th~Hyop~l~tions are 1i ke ly to be extirpated in the near future, resulting in theextTnctionofLRro­ pullans. iii ACKNOWLEDGEMENTS I am expressly grateful for patience and perceptive. advice during the course of this study to its completion. I am also especially grateful to Dr. Jerry Davis, my major advisor, and to Dr. Thomas Claflin, Dr. S. H. Sohmer, Dr. Thomas Weeks, and Dr. Weinzierl for their advice and .guidancewhiJe serving on my thesis conmittee. Mr. Mark Heitlinger and the Nature' Conservancy of Minnesota is gratefully acknowledged for their interest in and funding of this study. Each of the following individuals have made invaluable contributions to this study for which I am most grateful: Dr. Daniel Crawford, for his advice. and preliminary investigations in the chemotaxonomy of Erythronium; Dr. H~gh Iltis, for introducing me to the problem; Dr. Thomas Morley, for his advice and suggestions concerning Erythronium propullans; Mr. Orwin Rustad, for assisting me in locating populations ".+ c .....O/-h .... " ... and S. W. Batra, Dr. Lloyd Knutson,A. Menk~, and G. of the I sect Identification and Beneficial Insect JntrOcIuction Institute, Belts­ ville Agricultural Research Center, for their time and effort ih the identification of insects. Bruce Biltgen, Steve Swanson, and Diane Kell are given a special thanks for all of the help they have given me with this thesis during ~he past three years. iv TABLE OF CONTENTS PAGE LIST OF TABLES .. ·....... LIST OF FI GURES viii INTRODUCTION. .. 1 LITERATURE REVIEW ... 2 Taxonomy .. •• ..... 41: ••• 2 Cytology . ; ••. ·5 Life Hi story .. 5 STUDY SITE DESCRIPTION .. .. .. 10 Regional Description . 10 toea1ity Descri pi ons .. 17 MATERIALS AND METHODS .... .. .. .. 25 Morpho logy ... 25 Cytology . '. ........ 25 Demographics and Phenology . 26 ·Pollen and Ovule Production. 26 Pollen-Ovule Ratios ..... 27 Vectors of Pollen Transfer. 27 .~r~§cijrJ9 ~ tllciJ §!s 27 RESULTS ...•• .. 33 MOrphology •.......••• 33 Cyto logy.. .•...•••• Demog'raphics and Phenology ... Po 11 en and Ovu 1e Production. •. 44 Pollen-Ovule Ratios ....•. Observations of Floral Biology ...• Observations on Pollination Biology. Breeding Studies . 55 Seed Di spersa1 . • ...... DISCUSSION . 74 ""MQrRll()~l(lgy,.,~,==c'.=,"=~L.. •.•••••• •• ••• '.. .•..• ••• 74 Demographics and Phenology . ·... .. .. 74 Reproductive Capacity.... .. 78 Vectors of Pollen Transfer •. 78 Breeding Studies ..... .' ...... .. .. 80 Evolutionary Relationships. ......... 82 Species Persistence. ••••••• ·... ... 83 CONCLUSIONS •••••••.••• 85 . ...... 06 .... .'" . .. .. ·XION3ddV La . ·031IJ 3~nlV~31Il "vi LIST·"OF TABLES PAGE TABLE 1. Characteristics comparing .... ~.~.-_ ... (from Ireland, 1975, and D;;-h"",+.-,,,,.. 1Qhh\- ~- - .• •• 6, 2. COlllpariSonamongsites of 5 morphological characters an- alyzed in E. al bidum ....•.... - .... 34 3. Comparison among sites of 5 morphological characters an- alyzed in E.. propullans .....•, . .. .. 35 4. Comparison of morphological characters between E.pro- pull ans, the intermediate sizeelass, andLalbidum. .... 36 5. Number of flowering and non-flowering individualsofE. propull ans within: a .25 m2 area,wtth.th.e.g.pproximat~=- il.reaeach respective clone occupies. ...•......... 43 6. Pollen and ovule production and pollen-ovule ratios of ,I. propull ans and I. a1bidum .......•. 45 7. Percent stainable, unstainable and micropollen in E. pro­ pl.lllans andLal bidum ... ~-~---;-;--~--; ••.··•..i=...·.. J"~~~_,A.:~"~".j:'rbb between the sites utilized in 1977 the occurrance of apomixis in I. pro- ... .. ... .. ..... .. 56 9. Comparison of data between the sitesutilizedin1977 and 1978 measuring the occurrance of self-fertilization in I. propullans and I. albidum... .•.•..• ... •.... .. 58 10. Summary of res from treatment to det~rmine apomixis and self-fertilization within f. propullans and I· albidum in 1977 and 1978 .•.•....•...••....•.• .. 59 11. Comparison of data between the sites util ized in 1977 and 1978 measuring the success of intraclonal crosses in m ~... E. a1bi-dum· ,·e ..• ..>:.•.... r.···.. =oc=......... .. 60 12. Summary of results from treatments to determine success of intraclonal crosses in I. propul1ans~~~~. albidum in 1977 and 1978 .. .. .....•..• •• . - - . .. 61 vii 13. Comparison of data between sites measuring thesucc.ess of interclonal crosses in I. propullans and I. albidum in 1977 and 1978. •.••••••• .• •.•••.•••• 63 14. Summary of results from treatments to determine success of interclonal crosses in I. propullans and I. albidum in 1977 and 1978. .. ••••..••.•.•.•••.•.• . 64 15. Comparison of data between thesitesuti.l.i.iedjli-~J9_IZ~ and 1978 measuring the success of jnterpopulational crosses in E. propullans and I. albidum .•.... " ..• •. 65 16. Summary of results from treatments to determine the success··ofinterpopulational crosses in I. propullans and E. albidum in 1977 and 1978 .•. ... 66 17. Comparison of data between sites measuring thesuc.cess of interspecific hybridization between I. propullans and E. albidum in 1977 and 1978 .....•.••. •. .•• .• 68 18. Summary of results from treatments to determine success of interspecific hybridization between I .. propullansand E. a1bi dum i n1977 and 1978...-.--.-.---.-.----.---.--.-----.-----.-.. •. 69 19. Comparison of data between sites measuring the natural ~evel of seed production in I. propullans and I. albidum ln 1977 and 1978. ....•. • ...•.•. .. •.•• .. 70 20. SUl1llTlary of results determining the natural level of fruit and seed set of I. pro(JUlTa:i1s-and-r.--allJldUffiI--plus the annual seed crop of each species. .....••.•• .• 71 21. Comparison between I. propullans, the intermediate size class and E~albidum for 6 morphological characters ... •. 75 22. Comparison in the morphology, ecology, and chromosome number of I.propullans, I. albidum, and I. mesochoreum . .. 76 23. Summary of results obtained from breeding experiments in I. propullans 'and I ..albidum. ...•..••••.•.• .• 81 v;;; LIST OF FIGURES FIGURE Range of distribution of E. albidum! E. mesochoreum! 1. andE. propullans .... -: ...•..
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