Phylogenetic Development of the Inflorescence and Generic Relationships in the Kobresiaceae Charles Louis Gilly Iowa State College

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Phylogenetic Development of the Inflorescence and Generic Relationships in the Kobresiaceae Charles Louis Gilly Iowa State College Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1950 Phylogenetic development of the inflorescence and generic relationships in the Kobresiaceae Charles Louis Gilly Iowa State College Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Zoology Commons Recommended Citation Gilly, Charles Louis, "Phylogenetic development of the inflorescence and generic relationships in the Kobresiaceae " (1950). Retrospective Theses and Dissertations. 13679. https://lib.dr.iastate.edu/rtd/13679 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 directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely 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 will indicate the deletion. Oversize materials (e.g., nnaps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overiaps. ProQuest Information and Learning 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0600 NOTE TO USERS This reproduction is the best copy available. IMI" L PHYLOGENETIC DEVELOPMENT OP THE INFLORESCENCE AND GENERIC RELATIONSHIPS IN THE KOBRESIACEAE by Charles Louis Gllly A Dissertation Submitted to the Graduate Faculty in partial Fulfillment of The Requirements for the Degree of DOCTOR OP PHILOSOPHY Major Subject: Taxonomy Approved: Signature was redacted for privacy. Signature was redacted for privacy. Head of Major D^artment Signature was redacted for privacy. Iowa State College 1950 UMI Number: DP12713 UMI ® UMI Microform DP12713 Copyright 2005 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Q K-V ^ TABLE OP CONTENTS List of Tables v List of Plates (Figures and Maps) vi INTRODUCTION 1 REVIEW AND DISCUSSION OF LITERATURE 5 Classification of the Cyperaceae into Tribes 6 Principal Characteristics of the Tribe Cariceae 10 Division of the Tribe Cariceae into Genera l6 Inflorescence Development in the Tribe Cariceae 21 Pertinent Characteristics of the Family Gramineae 28 Geographical Origin and Possible Migration Routes 33 Cytogenetical Aspects of Evolution in the Tribe Cariceae 31 MATERIALS AND METHODS 3^ PHYLOGENETIC CONSIDERATIONS 58 Terminology of Inflorescence and Flower 60 Nature and Arrangement of Inflorescence Units 70 Modifications affecting flower arrangement 72 Modifications affecting Inflorescence axes 79 Structure and Homologies of the Perlgynium 8S Modifications affecting position of prophylls 82 Modifications affecting vasculariza­ tion of prophylls 84 Modifications affecting shape and texture of prophylls 86 Structure of Flowers and Fruit 89 Hi Modifications affecting carpel and stigma number 90 Modifications affecting style and apex of fruit 92 Modifications affecting staminate flowers 96 Structural Similarities of the Gramineae and Cariceae 97 DEVELOPMENTAL TRENDS IN THE INFLORESCENCE 101 Inflorescence Characteristics and Phases 101 Inflorescence Type-groups 107 Relative Advancement of Type-groups 123 TAXONOMIC CONSIDERATIONS 128 Nomenclatural Phases of the Problem 128 Kobresiaceae, Familia nova. Equivalent to Tribe Cariceae 13O Arrangement of Species of the Kobresiaceae in Species-groups 13^ GEOGRAPHIC CONSIDERATIONS 172 Distribution of Type-groups with Respect to Geographical Regions I76 Relative Advancement with Respect to Phytogeographical Regions 18I Geographical Distribution of Special Inflor­ escence Phases 185 Endemism and Discontinuous Distributions in Relation to Geographical Regions 200 Distributional History of the Kobresiaceae 2O5 The center of origin 206 Primary migration tracts and centers of distribution 211 Secondary migration tracts and centers of distribution 216 Tertiary distribution centers and spec iat ion 221 INTERRELATIONSHIPS OP SPECIES-GROUPS 227 Morphological Bases of Relationship 228 Iv Geographical Bases of Relationship 231 Phylogenetic Charts 2^9 SUMtlARY AND CONCLUSIONS 255 ACKNOWLEDGMENTS 26l LITERATURE CITED 263 APPENDICES 277 A. List of Species Referable to Type-groups 277 B. Advancement Values of Type-groups 308 C. Type-groups in Relation to Geographical Regions 312 V List of Tables 1. Relationship between Bracts, Prophylls and Flowers in the Carlcean In­ florescence 62 2. Number of Species, Type-groups and Species-groups in which Phases of In­ florescence Characteristics are Represented 106 3. Distribution of Type-groups with Refer­ ence to Number of Species 10? 4. Distribution of Type-groups and Species in Advancement Value Classes 125 5. Range of Advancement Values for Certain Genera or groups of Genera in the Complex 126 6. A comparison of the Cyperaceae (sensu strictior), Kobresiaceae and Gramineae 133 7. Number of Type-groups and Species-groups, with a Comparison of Advancement Value and Relative Rank of Geographical Regions 178 8. Number of Type-groups, Range and Mean of Advancement Value, and Relative Rank for Eleven Geographical Regions with Primitive and Advanced Type-groups 184 9. Comparison of Type-groups of the Kobres­ iaceae with Genera and Families of Anglosperms having Discontinuous Distributions 203 10. Comparison of Endemic Type-groups of the Kobresiaceae with Endemic Genera and Families of Anglosperms 204 11. Distribution of Species-groups with Respect to Center of Origin and Primary Distri­ butional Areas 219 vi List of Plates (Figures and Maps) I. Inflorescence Structure in the Kobresiaceae — 1 (Figures 1—7) 65 II. Inflorescence Structure in the Kobresiaceae — 2 (Figures 8—26) 68 III. Phylogenetic Development from an Androgynon (Figure 27) 7^ IV. Inflorescence Structure in the Kobresiaceae -- 3 (Figures 28—53) 8l V, Fruit and Flower Structure in the Kobresiaceae and the Genus Coix (Gramineae) (Figures 65—69) 95 VI. Phylogenetic Advancement in the Kobresiaceae (Figures 70—80) IO3 VII. Geographical Distribution of Species-groups of the Kobresiaceae — 1 (Figures 81—83) 1^3 VIII. Geographical Distribution of Species-groups of the Kobresiaceae — 2 (Figures 8^—86) 148 IX. Geographical Distribution of Species-groups of the Kobresiaceae — 3 (Figures 87—89) 153 X. Geographical Distribution of Species-groups of the Kobresiaceae — 4 (Figures 90—92) 159 XI. Geographical Distribution of Species-groups of the Kobresiaceae — 5 (Figures 93—95) 165 XII. Geographical Distribution of Species-groups of the Kobresiaceae —6 (Figures 96—98) I69 vll XIII. Geographical Relationships in the Kobresiaceae — 1 (Figures 99—101) 173 XIV. Geographical Relationships in the Kobresiaceae — 2 (Figures 102—104) I80 XV. Geographical Distribution of Inflorescence Phases — 1 (Figures IO5—107) I88 XVI. Geographical Distribution of Inflorescence Phases — 2 (Figures 108—110) I92 XVII. Geographical Distribution of Inflorescence Phases — 3 (Figures 111—II3) I96 XVIII. Geographical Distribution of Inflorescence Phases — 4 (Figures 114—II6) 199 XIX. Distributional History of the Kobres­ iaceae — 1 (Figures 117—119) 210 XX. Distributional History of the Kobresiaceae — 2, and Interrelationships of Inflorescence-type Units (Figures 120 and 121) 223 XXI. Interrelationships of Inflorescence- type Units with Arrangements of Staininate and Pistillate Flowers (Figures 122—126) 233 XXII. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 1 (Figure 127) 235 XXIII. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 2 (Figure 128) 237 XXIV. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 3 (Figure 129) 239 vill XXV. Morphological and Cfeographical Rela­ tionships of Type-groups in Species-groups — 4 (Figure 13O) 2^1 XXVI. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 5 (Figure 131) 243 XXVII. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 5 (Figure I32) 245 XXVIII. Morphological and Geographical Rela­ tionships of Type-groups in Species-groups — 7 (Figure 133) XXIX. Phylogenetic Chart of Inflorescence Type-groups in the Kobresiaceae (Figure 134) 251 XXX. Phylogenetic Chart of Species-groups in the Kobresiaceae (Figure I35) 253 1 INTRODUCTION Carex L., together with the three generally recognized related genera, Kobresla Wllld., Schoenoxiphiutn Nees and Uncinla Pers., has long been regarded as the nucleus of the tribe Cariceae of the family Cyperaceae. Within this family the Cariceae is distinct, easily recognizable and somewhat anomalousJ the most
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