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Download The A TAXONOMIC REVISION OF THE GENUS PHILONOTIS FOR NORTH AMERICA, NORTH OF MEXICO by WILLIAM MILTON ZALES B. S. ed. Eastern Illinois University, 1966 M. S. Eastern Illinois University, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of Botany We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, 19 73 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of BOTANY The University of British Columbia Vancouver 8, Canada Date 1 May 1973 i ABSTRACT The genus Philonotis (Musci) is revised for North /America, north of Mexico, and exclusive of Greenland,, following the last revision for this region by Seville Flowers in 1935. Collections and field observations of all but two of the recognized taxa were made from a wide diversity of habitats throughout most of their geographic range in North America. Numerous collections were cultured in a uniform environment to determine the stability of taxonomically diagnostic morphological characters. The position of papillae on the leaf cells, the structure of marginal cells, and to some degree the leaf shape and leaf cell shape are stable characters that are not modified by the environment. All other morphological •characters examined are ecophenic and cannot be utilized to segregate taxa. The present revision recognizes seven species and two varieties; Philonotis glaucescens (Hornsch.) Broth., P. longiseta (Rich, in Michx.) Britt., P. sphaerocarpa (Hedw.) Brid., P. mavohioa (Hedw.) Brid., P. capillaris Lindb. ex Hartm., P. fontana (Hedw.) Brid. var. fontana, P. fontana var. amevicana (Dism.) Flow, ex Crum, P. fontana var. pumila (Turn.) Brid., and P. yezoana Besch. et Card, in Card, (new to North America). Chromosome counts of n = 6 for P. glaucescens and n = 12 for P. longiseta are new reports. Keys, descriptions, illustrations, and distribution maps are provided for identification of the species. TABLE OF CONTENTS Abstract, i Table of Contents, ii-ila List of Tables, Hi List of Figures, iv-ivd List of Maps, v Acknowledgements, vi I. Introduction, 1 II- Field Work and Herbarium Studies, 3 III. Nomenclatural History of North American Philonotis, 4 IV. Methods of Analyzing Morphological Diversity, 9 V. Morphological Analysis of Philonotis from Herbarium and Field Specimens, 10 A. Sporophyte, 10 B. Gametophyte, 11 VI. Experimental Analysis of Morphological Diversity in Philonotis, 21 A. History of Experimental Culture in Philonotis, 21 B. Methods and Materials, 22 C. Results, 2 3 1. Leaf size, 24 2. Leaf shape, 24 3. Leaf apex, 25 4. Leaf lamina, 25 5. Leaf margin, 29 •6. Costa, 31 7. Leaf arrangement, 32 8. Stem, 32 9. Rhizoids, 32 10. Propagula, 33 D. Conclusions, 33 VII. Species Concept, 39 VIII. Taxonomic Treatment, 40 A. Description of the Genus in North America, 40 B. Key to the North American Species of Philonotis, C. Explanation of Species Figures, 45 Philonotis glaucescens, 46 Philonotis longiseta, 52 Philonotis sphaerocarpa, 60 Philonotis marchica, 63 Philonotis capillaris, 69 Philonotis fontana var. fontana, 73 Philonotis fontana var. americana, 82 Philonotis fontana var. pumila, 86 Philonotis yezoana, 90 IX. Phytogeography of Philonotis, 93 X. Distribution Maps, 95 XI. Excluded Taxa, 104 XII. Synonyms, 107 XIII. Summary, 111 XIV. Exsiccati and Selected Specimens Examined, 114 XV. Literature Cited, 162 • • • LIST OF TABLES ±. North American Species of Philonotis, 6 II. Morphological Characters of Philonotis, 36 III. Descriptive Names Referring to Ecophenic Modifications, 38 LIST OF FIGURES Figure 1-3 Philonotis fontana sporophyte and calyptra, 12 4, 5 Philonotis fontana stem and seta cross section, 13 6* 7 Philonotis fontana peristome teeth, 14 8-15 Scanning electron micrographs of the spores of North American species of Philonotis , 15 16-25 Morphological variability in Philonotis fontana induced by seasonal growth, 18 26-33 Comparison of various ecophenic expressions of Philonotis fontana, 26 34, 35 Leaves of Philonotis glauoescens, 26 36-40 Types of lamina cells found in the North American species of Philonotis, 30 41-43 Types of leaf margin cells found in the North American species of Philonotis, 30 44, 45 Scanning electron micrographs of Philonotis fontana rhizoids produced on the same stem, 34 46-48 Comparison of leaf shape and leaf margin in Philonotis glauoescens 3 marchica, and sphaerooarpa, 49 49 Philonotis glauoescens, 51 50, 51 Mitotic chromosome configurations of Philonotis longiseta and Philonotis glauoescens, 55 52, 53 Comparison of leaf shape and median leaf cell shape of Philonotis marchica and Philonotis longiseta, 57 54 Philonotis longiseta, 59 55 Philonotis sphaerooarpa, 62 56, 57 Comparison of leaf shape, apex, and median leaf cells of Philonotis marchica and Philonotis capillaris, 67 58 Philonotis marchica, 68 59 Philonotis capillaris, 72 '60 Philonotis fontana var. fontana, 81 61 Philonotis fontana var. americana, 85 62 Philonotis fontana var. pumila, 89 63 Philonotis yezoana, 92 V LIST.OF DISTRIBUTION MAPS Philonotis glauoescens, 95 Philonotis longiseta, 96 Philonotis sphaerooarpa, 97 Philonotis marchica, 98 Philonotis capillaris, 99 Philonotis fontana var. fontana, 100 Philonotis fontana var. americana, 101 Philonotis fontana var. pumila, 102 Philonotis yezoana, 103 vi ACKNOWLEDGEMENTS I wish to express gratitude to my graduate advisor Dr. W. B. Schofield for his influence and direction in demonstrating the value of thorough literature reviews, persistent field work, and for reading this manuscript throughout its production. I am also indebted to the members of the Botany Department of the University of British Columbia who in numerous ways have assisted and encouraged my progress towards the completion of this research. I wish to thank the curators of the herbaria listed on page 3 for their time and cooperation in sending specimens on loan for this study. Acknowledgement is made of financial aid from the National Research Council of Canada through grants to Dr. W. B. Schofield for summer field expenses. I am especially thankful for the patience and sympathetic understanding of my wife Dotty, and for the encouragement and assistance she has given during this project. 1 I. INTRODUCTION The well defined moss genus Philonotis of the family Bartramiaceae contains approximately 182 highly variable species that occur in moist or wet habitats throughout the world. In North America (exclusive of Greenland and Mexico) members of the genus are recognized by loose or dense tufted plants usually clothed in abundant tomentum, with short slender (1 cm high) stems varying to robust branched plants as large as 10 cm high that often possess whorled innovations and propaguliferous shoots or both. The highly variable leaves are ovate or triangular-lanceolate with an obtuse or acute apex and a long stout costa. The cells of the lamina are variously rectangular with prominent papillae and the "margins are toothed with single or double teeth and usually rolled. The perigonia are discoid (rosette shaped) or gemmiform (bud shaped) and all but P. longiseta are dioicous. The sporophyte is globose, inclined, and longitudinally furrowed on a long seta with a double series of sixteen peristome teeth. Philonotis can tolerate a wide range of ecological conditions although it is unusual on living trees and rare on rotten wood. The wide ecological range is paralleled by a wide degree of morphological plasticity. This polymorphism has resulted in the description of many subspecific taxa. In the North American flora alone 99 names are involved. The descriptions accompanying some taxa match a number of specimens. Other diagnoses agree with the descriptions of 2 several different taxa, and the remainder fit no available description. In the present treatment seven distinct species and two varieties are recognized. Careful scrutiny of the available literature exposes the fact that many errors and mis• understandings have been perpetuated from one author's work to another. Even more disheartening is the realization that the identifications of previous monographers of portions of the genus (Dismier, Flowers, Kindberg intev alia) do not properly match their own keys and descriptions. The present research, therefore, attempts to eliminate the ambiguity that plagues the genus in North America and to clarify the species concept. The purpose of the present treatise is to define the morphological variability of the genus Philonotis, to circumscribe its taxa, and to provide keys, illustrations, and maps to aid in their identification. To accomplish these goals numerous collections cultured in a uniform environment were used to assess the natural variability of critical morphological characters within the species. Such cultures should reveal whether these characters are stable or whether they are variable (ecophenic) depending on various ecological parameters. An ecophene is defined as a plant or group of plants that show a changed morphology induced by an altered environment (ecologically modified phenotype). II. FIELD WORK AND
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