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Taxonomic revision of the genus Philonotis for North America, north of Mexico Zales, William Milton 1973

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A TAXONOMIC REVISION OF THE GENUS PHILONOTIS FOR NORTH AMERICA, NORTH OF MEXICO by WILLIAM MILTON ZALES B. S. ed. Eastern I l l i n o i s University, 1966 M. S. Eastern I l l i n o i s 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 t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, 19 73 In presenting t h i s t h e s i s in p a r t i a l f u l f i l m e n t o f the requirements for an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I f u r t h e r agree t h a t permission f o r e x t e n s i v e copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r e p r e s e n t a t i v e s . I t i s understood that copying or p u b l i c a t i o n of t h i s t h e s i s for f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission. Department of BOTANY The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date 1 May 1973 i ABSTRACT The genus Philonotis (Musci) i s revised for North /America, north of Mexico, and exclusive of Greenland,, following the l a s t r e v i s i o n for t h i s region by S e v i l l e Flowers i n 1935. Collections and f i e l d observations of a l l but two of the recognized taxa were made from a wide d i v e r s i t y of habitats throughout most of t h e i r geographic range i n North America. Numerous c o l l e c t i o n s were cultured i n a uniform environment to determine the s t a b i l i t y of taxonomically diagnostic morphological characters. The p o s i t i o n of p a p i l l a e on the leaf c e l l s , the structure of marginal c e l l s , and to some degree the l e a f shape and l e a f c e l l shape are stable characters that are not modified by the environment. A l l other morphological •characters examined are ecophenic and cannot be u t i l i z e d to segregate taxa. The present r e v i s i o n recognizes seven species and two v a r i e t i e s ; Philonotis glaucescens (Hornsch.) Broth., P. longiseta (Rich, in Michx.) B r i t t . , P. sphaerocarpa (Hedw.) Brid . , P. mavohioa (Hedw.) B r i d . , P. capillaris Lindb. ex Hartm., P. fontana (Hedw.) B r i d . var. fontana, P. fontana var. amevicana (Dism.) Flow, ex Crum, P. fontana var. pumila (Turn.) Bri d . , 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, i l l u s t r a t i o n s , and d i s t r i b u t i o n maps are provided for i d e n t i f i c a t i o n of the species. TABLE OF CONTENTS Abstract, i Table of Contents, i i - i l a L i s t of Tables, Hi L i s t of Figures, iv-ivd L i s t of Maps, v Acknowledgements, vi I. Introduction, 1 II- F i e l d Work and Herbarium Studies, 3 I I I . Nomenclatural History of North American Philonotis, 4 IV. Methods of Analyzing Morphological D i v e r s i t y , 9 V. Morphological Analysis of Philonotis from Herbarium and F i e l d Specimens, 10 A. Sporophyte, 10 B. Gametophyte, 11 VI. Experimental Analysis of Morphological Div e r s i t y i n Philonotis, 21 A. History of Experimental Culture i n Philonotis, 21 B. Methods and Materials, 22 C. Results, 2 3 1. Leaf s i z e , 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 i n 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. D i s t r i b u t i o n Maps, 95 XI. Excluded Taxa, 104 XII. Synonyms, 107 XIII. Summary, 111 XIV. E x s i c c a t i and Selected Specimens Examined, 114 XV. L i t e r a t u r e 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 v a r i a b i l i t y i n 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 c e l l s found i n the North American species of Philonotis, 30 41-43 Types of l e a f margin c e l l s found i n 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 l e a f shape and leaf margin i n Philonotis glauoescens 3 marchica, and sphaerooarpa, 49 49 Philonotis glauoescens, 51 50, 51 M i t o t i c chromosome configurations of Philonotis longiseta and Philonotis glauoescens, 55 52, 53 Comparison of l e a f shape and median le a f c e l l shape of Philonotis marchica and Philonotis longiseta, 57 54 Philonotis longiseta, 59 55 Philonotis sphaerooarpa, 62 56, 57 Comparison of l e a f shape, apex, and median l e a f c e l l s 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 d i r e c t i o n i n demonstrating the value of thorough l i t e r a t u r e reviews, persistent f i e l d work, and for reading t h i s manuscript throughout i t s production. I am also indebted to the members of the Botany Department of the University of B r i t i s h Columbia who i n numerous ways have assisted and encouraged my progress towards the completion of t h i s research. I wish to thank the curators of the herbaria l i s t e d on page 3 for t h e i r time and cooperation i n sending specimens on loan for t h i s study. Acknowledgement i s made of f i n a n c i a l aid from the National Research Council of Canada through grants to Dr. W. B. Schofield for summer f i e l d expenses. I am e s p e c i a l l y thankful for the patience and sympathetic understanding of my wife Dotty, and for the encouragement and assistance she has given during t h i s project. 1 I. INTRODUCTION The well defined moss genus Philonotis of the family Bartramiaceae contains approximately 182 highly variable species that occur i n 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 i n 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 c e l l s of the lamina are variously rectangular with prominent p a p i l l a e and the "margins are toothed with sing l e or double teeth and usually r o l l e d . The perigonia are d i s c o i d (rosette shaped) or gemmiform (bud shaped) and a l l but P. longiseta are dioicous. The sporophyte i s globose, i n c l i n e d , and l o n g i t u d i n a l l y furrowed on a long seta with a double series of sixteen peristome teeth. Philonotis can tolera t e a wide range of e c o l o g i c a l conditions although i t i s unusual on l i v i n g trees and rare on rotten wood. The wide e c o l o g i c a l range i s p a r a l l e l e d by a wide degree of morphological p l a s t i c i t y . This polymorphism has resulted i n the description of many subspecific taxa. In the North American f l o r a alone 99 names are involved. The descriptions accompanying some taxa match a number of specimens. Other diagnoses agree with the descriptions of 2 several d i f f e r e n t taxa, and the remainder f i t no available d e s c r i p t i o n . In the present treatment seven d i s t i n c t species and two v a r i e t i e s are recognized. Careful scrutiny of the avail a b l e l i t e r a t u r e exposes the fact that many errors and mis-understandings have been perpetuated from one author's work to another. Even more disheartening i s the r e a l i z a t i o n that the i d e n t i f i c a t i o n s of previous monographers of portions of the genus (Dismier, Flowers, Kindberg intev alia) do not properly match t h e i r own keys and descriptions. The present research, therefore, attempts to eliminate the ambiguity that plagues the genus i n North America and to c l a r i f y the species concept. The purpose of the present t r e a t i s e i s to define the morphological v a r i a b i l i t y of the genus Philonotis, to circumscribe i t s taxa, and to provide keys, i l l u s t r a t i o n s , and maps to aid i n t h e i r i d e n t i f i c a t i o n . To accomplish these goals numerous c o l l e c t i o n s cultured i n a uniform environment were used to assess the natural v a r i a b i l i t y of c r i t i c a l morphological characters within the species. Such cultures should reveal whether these characters are stable or whether they are variable (ecophenic) depending on various e c o l o g i c a l parameters. An ecophene i s defined as a plant or group of plants that show a changed morphology induced by an altered environment (ec o l o g i c a l l y modified phenotype). II. FIELD WORK AND HERBARIUM STUDIES During the summers of 19 71 and 19 72 approximately 330 specimens of Philonotis were c o l l e c t e d i n southeastern, eastern, and western North America. At these times general f i e l d observations were recorded to obtain an appreciation of the ecol o g i c a l amplitude of each species c o l l e c t e d . A l l but three (P. yezoana, P. sphaevoaarpa, and P. fontana var. pumila) of the nine taxa have been cultured i n a uniform environment chamber, and the l a t t e r two have not been observed in the f i e l d . Numerous specimens, including a l l available type specimens, have been examined from various herbaria i n North America, Europe, and Asia. Over 10,500 specimens were borrowed from the following herbaria: (standard abbreviations according to Lanjouw & Sta f l e u 1964) B, BM, BRNM, C, CAN, COLO, CU, DPU, DUKE, F, FLAS, FSU, G, GL, GRI, GRO, H, IA, L, LAF, LD, MANCH, MICH, MO, NFLD, NO, NY, OP, PC, SMS,,SMU, S-PA, TENN, TEX, TRTC, UAC, UBC, UC, UPSV, US, UT, WIS, WJC, and To t t o r i University, Japan (TOT). 4 III. N 0 F 1 E N C L A T U R A L HISTORY OF NORTH AMERICAN PHILONOTIS Three species that are now included i n the genus Philonotis {fontanum, marchicum, and sphaericarpon) were described by Hedwig i n 1801 as belonging to the large and i n c l u s i v e genus Mnium. Hedwig also described two species of Bartramia but f a i l e d to draw any relationships at the family l e v e l between these taxa and the three species of "Mnium", probably due to t h e i r difference i n sexuality. This i s understandable because of the acceptance of Linnaeus 1 dependence on sexual characters for the c l a s s i f i c a t i o n of flowering plants. B r i d e l (1826) described the genus Philonotis separating i t from Mnium and included i n i t P. fontana with v a r i e t i e s alpina and falcata, P. marchica, and P. sphaerocarpa. At t h i s time no family relationships were established with the genus Bartramia. In the important European work Bryologia Europaea, (Bruch et al. 1842) many new families and genera were described. Such large and unnatural Hedwigian genera as Mnium and Hypnum were divided into several more homogeneous genera. The family Bartramiaceae was also established and included Oreas, (now included i n the Dicranaceae), Bartramidula, and Bartramia. B r i d e l ' s species of Philonotis, however, were included i n Hedwig's o r i g i n a l , but now enlarged genus, Bartramia. Schimper (1855) i n the Corgllarium to Bryologia Europaea reinstated B r i d e l ' s genus Philonotis with the addition of two subgeneric categories, Philonotula, with gemmiform perigonia, and Philonotis, with d i s c o i d perigonia. Except for authors who have described new taxa within the genus, Schimper's treatment i s where the generic concept has remained u n t i l the present. Table I l i s t s the taxa included i n catalogues of North American mosses by various authors. These are arranged i n chronological order. 6 TABLE I. NORTH AMERICAN SPECIES OF PHILONOTIS Rail & Hervey 1880 Catalogue of North American Musci Philonotis marchica B r i d . {muehlenbergii Schwaegr.) fontana L. var. falcata (no authority cited) radicalis Beauv. var. porteri Aust. tenella Hedw. Lesquereux & James 1884 Manual of the Mosses of North America Philonotis muehlenbergii B r i d . var. tenella B r i d . macounii Lesq. et James sp. nov. fontana B r i d . var. alpina B r i d . var. falcata B r i d . calcarea Schimp. mohriana (C. Mull.) Jaeg. Kindberg 1897 European and North American Bryineae Philonotis marchica W i l l d . muehlenbergii B r i d . macounii Lesq. et James arnellii Husn. vancouveriensis Kindb. sp. nov. glabriuscula Kindb. calcarea Br. Eur. var. caespitosa Wils. alpicola Jur. acutiflora Kindb. fontana L. var. seriata Mitt. mohrii C. Mull. adpressa Ferg. radicalis Beauv. tenella C. Mull. pumila Kindb. sp. nov. Dismier 1910 Philonotis de I'Amerique Philonotis radicalis (P. B.) Br i d . gracillima Angstr. sphaerooarpa (Sw.) Bri d . ssp. tenella (C. Mull.) Jaeg. var. terrestris Dism. var. nov. 7 marchica (Willd.) B r i d . seriata (Mitt.) Lindb. ssp. americana Dism. ssp. nov. var. torquata (Ren. et Geh. ex Geh.) Dism. var. graciles cens Dism. var. nov. fontana B r i d . var. heterophylla Card, et Ther. ssp. tomentella Mol. var. borealis (Hag.) Loeske var. compacta Dism. var. heterophylla Dism. var. nov. caespitosa Wils. var. compacta Dism. var. nov. var. laxa (Warnst.) Loeske et Warnst. var. adpressa Dism. var. heterophylla Dism. var. nov. ssp. fallax Dism. ssp. nov. oapillaris Lindb. S. Flowers 1935 i n Grout, Moss Flora of North America Philonotis glauoescens (Hornsch.) Paris f. laxa Flow. /. nov. var. terrestris (Dism.) Flow. comb. nov. var. brevifolia (Dism.) Flow. comb. nov. gracillima Angstr. sphaericarpa (Sw.) Br i d . var. terrestris Dism. uncinata (Schwaegr.) B r i d . longiseta (Rich.) E. G. B r i t t . f. propagulicaulis Flow. f. polygama Flow. /. nov. var. porteri (Aust.) comb. nov. marchica (Willd.) B r i d . muehlenbergii (Schwaegr.) B r i d . oapillaris Lindb. caespitosa Wils. var. compacta Dism. var. adpressa Dism. var. laxa (Warnst.) Loeske fontana B r i d . f. dimorphophylla Flow. /. nov. var. heterophylla Card. & Ther. var. seriata B r e i d l . f. occidentalis Flow. /. nov. var. pumila B r i d . f. longifolia Flow. nov. f. dimorphophylla Flow. /. nov. var. falcata B r i d . var. adpressa (Ferg.) Limpr. var. borealis Hag. 8 var. laxa Flow. var. nov. f. tenuis Flow. f. nov.. amerioana Dism. f. taxa Flow. /. nov. f. dimorphophylla Flow. /. nov. var. torquata (Ren. et Geh.) Dism. var. graciles cens Dism. oalcarea f. occidentalis Flow. /. nov. Crum, Steere, & Anderson 1965 A List of the Mosses of North America Philonotis amerioana Dism. var. gracilescens Dism. var. torquata (Ren. et Geh. ex Geh.) Dism. caespitosa Jur. var. adpressa Dism. var. ampliretis (Dix.) Crum, Steere, et Anderson var. compacta Dism. oalcarea (B. S. G.) Schimp. capillaris Lindb. fontana (Hedw.) Bri d . var. adpressa (Ferg. ex Hunt) Limpr. var. borealis I. Hag. var. falcata B r i d . var. heterophylla Card, et Ther. var. laxa Flow. var. pumila B r i d . var. seriata (Mitt.) B r e i d l . glaucescens (Hornsch.) Par. var. terrestris (Dism.) Flow. gracillima Angstr. longiseta (Michx.) B r i t t . var. porteri (Aust.) Flow. marchica (Hedw.) Bri d . muehlenbergii (Schwaegr.) Brid. sphaerocarpa (Hedw.) Brid. uncinata (Schwaegr.) B r i d . 9 IV. METHODS OF ANALYZING MORPHOLOGICAL DIVERSITY Morphological d i v e r s i t y may be studied i n several ways. (i) Perennial plants l i k e Philonotis may show v a r i a t i o n due to seasonal changes i n the environment. This v a r i a t i o n i s e a s i l y detected by examining one year's growth produced on a plant. Hi) Examination of several plants from a clo n a l population may show v a r i a t i o n induced by competition for water, space, and minerals, or d i f f e r e n t degrees of exposure to sunlight. {Hi) C r i t i c a l examination of large series of c o l l e c t i o n s from throughout the entire range of a taxon may demonstrate that v a r i a t i o n i s e i t h e r a continuum connecting several taxa with many intermediate forms or that there are d e f i n i t e groups of variants that may be recognized taxonomically. This i s c l e a r l y the most important consideration i n t r a d i t i o n a l or alpha taxonomy. Unfortunately many taxa of Philonotis have been described from a single specimen with no allowance given for v a r i a b i l i t y . (iv) Mor-phological v a r i a b i l i t y that i s suspected to be environmentally induced can be tested by c u l t i v a t i n g morphologically d i s t i n c t specimens from numerous locations under uniform environmental conditions and comparing the new growth that develops. . A l l four methods of analysis are considered herein, however the experimental results have proven to be the most r e l i a b l e for revealing morphological d i v e r s i t y and i t s ' causes i n Philonotis. 10 V. MORPHOLOGICAL ANALYSIS OF PHILONOTIS FROM HERBARIUM AND FIELD SPECIMENS A. SPOROPHYTE Although there i s no a priori reason why the sporophytic c h a r a c t e r i s t i c s of a moss should be less p l a s t i c or variable than the vegetative haploid gametophyte, the sporophyte i s the most stable phenotypic expression of Philonotis. The i n i t i a t i o n and production of gametangia, transfer of sperms, f e r t i l i z a t i o n of the egg, and subsequent maturation of the sporophyte requires very s p e c i f i c environmental conditions i n order to be successful. Unfortunately this narrow e c o l o g i c a l tolerance d r a s t i c a l l y reduces the number of specimens c o l l e c t e d with mature sporophytes. Not only are the sporophytes of l i t t l e value as diagnostic characters due to t h e i r s c a r c i t y but the sporophyte i s also morphologically uniform throughout the genus. This viewpoint i s also held by Dismier (1907a) . The seta i s green when young but becomes brown when mature and i s twisted throughout when dry. Seta length i s variable (1-6 cm) but generally longer in the larger specimens of P. fontana. The anatomy i s s i m i l a r i n a l l North American species ( f i g . 5). The capsule i s spherical and l i g h t green while developing ( f i g . 2). I t becomes brown, deeply furrowed l o n g i t u d i n a l l y , and constricted about the middle when mature and empty ( f i g . 3). The capsule i s held obliquely and the peristome i s positioned further o f f the v e r t i c a l axis of the seta ( f i g . 2, 3). The exothecial c e l l s are i r r e g u l a r l y 11 rectangular, becoming i r r e g u l a r l y cuboidal at the base where the stomata are located. Forty to 275 stomata have been counted on various capsules however the maximum to minimum ranges of a l l North American species overlap between 90 and 100 stomata per capsule. The c e l l walls are thicker on the ridges than i n the furrows. The operculum i s round-convex with a shallow apiculus i n a l l species examined ( f i g . 2). The peristome consists of two c i r c l e s of sixteen teeth each that are s i m i l a r for a l l species ( f i g . 6, 7). The chromosome number of Philonotis i s constantly n = 6 i n a l l species reported except for P. longiseta and P. socia which are n = 12. (See discussion under P. longiseta.) B. G A M E T O P H Y T E Spore morphology i n Philonotis does not provide any consistent diagnostic features. Spore size i s variable depending on the siz e and maturity of the capsules they were produced i n . C r i t i c a l examination of spores with the scanning electron microscope (SEM) has revealed that the North American species are a l l s i m i l a r l y p a p i l l o s e and that the degree of p a p i l l o s i t y i s variable ( f i g . 8-15). The small, smooth, and cucullate calyptra i s shed very early i n sporophyte development. The few calyptras examined seemed to be i d e n t i c a l i n several d i f f e r e n t species ( f i g . 1). Their s c a r c i t y and s i m i l a r i t y make them useless for taxonomic use within the genus Philonotis. The p e r i g o n i a l and p e r i c h a e t i a l leaves surrounding the gametangia have h i s t o r i c a l l y been employed as diagnostic Figures 1-3. Philonotis fontana sporophyte & calyptra -1. Calyptra -2. Fresh mature capsule with operculum -3. Dry empty capsule 12 1 mm F i g u r e s 4, 5. Philonotis fontana stem & s e t a c r o s s s e c t i o n -4. Stem c r o s s s e c t i o n -5. S e t a c r o s s s e c t i o n Figures 6,7. Philonotis -6. Portion of -7. Portion of fontana peristome teeth endostome exostome 14 Figures 8-15. Scanning electron micrographs of the spores of North American species of Philonotis -8. P. glauoes oens -9. P. longiseta -10. P. sphaerocarpa -11. P. marchica -12. P. capillaris -13. P. fontana var. fontana -14. P. fontana var. amerioana -15. P. fontana var. pumila 16 characters i n Philonotis to an unusually great extent. The two largest sections of Philonotis are organized according to the arrangement of leaves forming the perigonia: section Philonotis possesses d i s c o i d perigonia, and includes those species with sguarrose-geniculate leaves forming a f l a t rosette; whereas section Philonotula has gemmiform perigonia, and includes those with erect leaves forming a bud. At the s e c t i o n a l l e v e l t h i s character (the arrangement of p e r i g o n i a l leaves) i s consistent, but at the s p e c i f i c l e v e l the perigonia are of s l i g h t value for two basic reasons. F i r s t , specimens are not always c o l l e c t e d i n a sexual condition, and second these leaves show an extreme polymorphism within each general type. Janzen (1921), Loeske (1906), and Monkemeyer (1927) have studied the v a r i a b i l i t y of p e r i g o n i a l and p e r i c h a e t i a l leaves of Philonotis i n great d e t a i l and also agree that they are of l i t t l e taxonomic value i n d i s t i n g u i s h i n g species. The flask shaped archegonia, oblong antheridia, and the numerous elongate club shaped paraphases, which are s i m i l a r to most other bryophytes, are uniform throughout the North American species. Polymorphism as observed i n the vegetative leaves of Philonotis i s also extreme and has resulted i n the proposal of many superfluous names. The s i z e , color, contour, and to some degree the shape of the leaves, and t h e i r p o s i t i o n and arrangement on the stem are a l l subject to modification by the environment. Meusel (1935), and i n greater d e t a i l Hagerup (1935), described and i l l u s t r a t e d ( f i g . 16-25) the periodic or seasonal growth of P. fontana from f i e l d observations. They both remarked on the production of gametangia at the base of several innovations or shoots so c h a r a c t e r i s t i c of the genus. Hagerup (19 35) continued to explain the presence of two kinds of annual shoots: spring shoots with short, blunt, c l o s e l y appressed leaves ( f i g . 17, 23), and summer shoots with lanceolate, somewhat falcate leaves ( f i g . 20, 22). He made no taxonomic statements or suggestions even though P. fontana var. heterophylla Card, et Ther. corresponds to f i g . 18, and P. fontana f. dimorphophylla Flow, corresponds to f i g . 20. These two subspecific taxa are r e c i p r o c a l dimorphic conditions produced i n response to seasonal change, and should not be accorded separate taxonomic status. I f an enti r e t u f t or sod i s examined rather than a single plant, a better appreciation i s gained of the polymorphism within a taxon. I t may be assumed that the plants of a t u f t have the same genotype as they are often the product of vegetative reproduction. I f the t u f t i s f e r t i l e and reproducing sexually we may usually assume that the parents are of the same gene pool and that a l l of the ind i v i d u a l s belong to the same taxon. Plants i n the center of a t u f t w i l l generally be more robust, more tomentose, and i n some species posses larger, f a l c a t e , or p l i c a t e leaves with r o l l e d margins. Plants at the edge of a population are depauperate, lax, l i g h t e r i n color, less erect, often Figures 16-25. Morphological v a r i a b i l i t y i n Philonotis fontana induced by seasonal growth -17&23. Spring shoot and leaf -20&22. Summer growth on the same shoot and le a f 24. One innovation (branch) of a male plant becoming f e r t i l e i n the second year 25. Mature sporophyte produced on a female plant during the t h i r d year, and a young sporophyte developing at the end of the same year's growth After Hagerup (19 35 p. 8&10) 18 19 e t i o l a t e d , and with f l a t appressed leaves. These extremes i n a population correspond to described v a r i e t i e s , forms, and even other species, such as adpressa, falcata, taxa, graailescens3 eaespitosa, and oompaota. I t i s noteworthy that these taxa are rarely c o l l e c t e d with sporophytes. Compared to the narrow e c o l o g i c a l requirements necessary to produce mature sporophytes, the gametophyte can t o l e r a t e a wide range of environmental conditions. Those conditions that favor sporophyte production w i l l also produce the he a l t h i e s t gametophytes whereas other environmental conditions w i l l produce ent i r e populations that correspond to these s t e r i l e v a r i e t i e s , forms, and "species". Polymorphism i s even more confusing when plants from widely separated populations are compared. Then the assumption that these taxa actually belong to a single species i s even more d i f f i c u l t to support unless numerous series of c o l l e c t i o n s are examined, and the described taxa are shown to be connected by a complete l i n e of intermediate and intergrading forms. After examining a l l available North American and many extra-North American specimens, i t seems plausible to agree with Flowers (s. a.). " I t i s time to give up the pretense that cer t a i n segments of t h i s array of plants represents discrete genetical groups and face up to the fact that we are dealing with a highly involved s i t u a t i o n where the i n t e r a c t i o n of 20 heredity and environment i s producing a maze of forms stemming from e s s e n t i a l l y the same stock." VI. EXPERIMENTAL ANALYSIS OF MORPHOLOGICAL DIVERSITY IN PHILONOTIS A. HISTORY OF EXPERIMENTAL CULTURE OF PHILONOTIS Experimental studies using bryophytes i n culture have been concerned almost exclusively with the physiology of a very few common plants l i k e Funaria hygrometvica or Marohantia polymorpha. H . Buch (1922) conducted extensive studies on the hepatic family Scapaniaceae i n uniform environment chambers and found that water and l i g h t influenced the plant's morphology considerably. More recently Lodge (1960) cultured Drepanooladus fluitans and D. exannulatus and found that the shape and arrangement of the leaves and the length of the costa are highly variable depending mostly on the amount of moisture and l i g h t . Davy de V i r v i l l e (1927) studied the action of the environment on several mosses and found that Philonotis fontana i n a saturated atmosphere produces many rhiz o i d s , has shorter leaves with thinner c e l l walls, smaller teeth, and fewer chloroplasts. When the plant i s grown submerged i t has l i g h t green stems 2-4 cm long with few rh i z o i d s , variable leaves e x h i b i t i n g an attenuated apex, thin c e l l walls, and large chloroplasts. Zastrow (19 34) grew P. fontana i n l i q u i d culture and found that i t tolerated a wide pH range, developed long stems with widely spaced erect leaves that had f l a t margins, a thi n 22 percurrent costa without a central strand, and lax c e l l s , but retained t h e i r o r i g i n a l shape. Extensive research has been conducted by many workers on spore v i a b i l i t y , mode of germination, and the ph y s i o l o g i c a l e f f e c t s that control or influence spore germination. Only Matteri (19 70) has studied members of the Bartramiaceae and she found that three species of Philonotis develop the t y p i c a l heterotrichous type of protonema and no intrageneric differences were present. No one has cultured Philonotis for the purpose of studying the v a l i d i t y of i t s taxonomic characters, although several authors have suggested that t h i s type of experimental research might be f r u i t f u l . (Nyholm 1960, Florschiitz 1964, Bowering 1966). The present research shows that some morphological features of the genus Philonotis are stable, while others that have t r a d i t i o n a l l y been thought to be important are too variable to be of taxonomic value. B . M E T H O D S A N D M A T E R I A L S A uniform environment chamber was constructed for the purpose of cu l t u r i n g large c o l l e c t i o n s of Philonotis from diverse populations. The chamber measured 48 X 96 inches and had a capacity of approximately 270 p e t r i dishes. The cultures were illuminated by eight 36 inch cool white fluorescent and eighteen 7 watt incandescent white bulbs suspended 20 inches above the surface of the dishes. A t o t a l i l l u m i n a t i o n of approximately 200 foot candles was produced. The chamber was automatically adjusted to maintain an 23 18/6 hour day/night cycle. The temperature varied with that of the ambient a i r and increased approximately 10°F during each day cycle. High humidity was maintained by frequent watering i n the dishes with 50% Hoagland's solution (Hoagland & Arnon 19 38). Small populations of plants, approximately 5X5 cm, from various habitats and various locations throughout North America were placed on wet paper toweling i n standard p l a s t i c p e t r i dishes. After several weeks the new growth was transferred to s t e r i l e Hoagland's solution s o l i d i f i e d with 2% agar. This process was continued u n t i l obvious fungal and a l g a l contamination was eliminated. A l l observations, measurements, and i l l u s t r a t i o n s were made from s l i d e s prepared by f i r s t relaxing the plants i n b o i l i n g water. The plants were then placed i n lacto-phenol and the leaves were removed from the stem. Lacto-phenol i s composed of equal parts of phenol, l a c t i c acid, g l y c e r i n , and d i s t i l l e d water (Johansen 1940). The lacto-phenol was drained and the specimens were made permanent i n Hoyer's mounting medium (Anderson 1954) . C. RESULTS In culture a l l of the taxa show a high degree of morphological s t a b i l i t y , although t h e i r appearance may be d r a s t i c a l l y d i f f e r e n t from the o r i g i n a l c o l l e c t i o n i n nature. This demonstrates that a s p e c i f i c set of environmental conditions w i l l produce a s p e c i f i c set of morphological responses. The comparison of s p e c i f i c characters as they 24 appear i n nature to t h e i r appearance i n culture can be used to assess t h e i r diagnostic value. Comparison of several morphologically d i s t i n c t plants from various habitats i n nature with t h e i r new growth produced i n a uniform environment has shown that t h e i r d i s t i n c t i v e differences are no longer present, and that they have become morphologically i d e n t i c a l . These " d i s t i n c t i v e " differences can then be assumed to be ecophenic or environmentally induced, and the plants can be assumed to belong to the same taxon. Diagnostic characters of Philonotis are analyzed and evaluated i n d i v i d u a l l y based on the results obtained from culture experiments. 1. LEAF SIZE Leaf size i s one of the most variable c h a r a c t e r i s t i c s of Philonotis. In general the leaves found in nature are much larger than the ones that develop i n culture, although plants with small leaves are often c o l l e c t e d . Unless f e r t i l e plants are c o l l e c t e d , l e a f s i z e i s misleading and other characters must be sought for analysis. 2. LEAF SHAPE The general leaf shapes are consistent i n some respects and variable i n others. Polymorphism, produced by d i f f e r e n t i a l seasonal growth, observed i n some species i n nature, i s l o s t i n culture; dimorphic and heteromorphic forms ( f i g . 16-25) produce only one kind of leaf i n a uniform environment. Plants with falcate leaves i n nature produce only s t r a i g h t leaves i n culture ( f i g . 28, 30). Very broadly ovate-lanceolate leaves become lanceolate i n culture ( f i g . 27, 29, 31). Species with triangular-lanceolate leaves i n general are stable and can be distinguished from species with ovate-lanceolate leaves. From the analysis of culture studies, leaf shape i s a suitable character only when used i n combination with other characters. 3 , LEAF APEX The l e a f apex remains the same i n culture as i n the f i e l d i n a l l taxa except P. glaucescens. Plants of P. glaucescens with an acuminate apex and percurrent costa i n the f i e l d w i l l develop a rounded apex with a shorter costa i n culture ( f i g . 34). Plants with a rounded apex i n nature maintain t h i s condition i n culture but when transplanted to a suitable environment i n the f i e l d develop an acuminate apex ( f i g . 35) . When these plants from the f i e l d were returned to the culture chamber they again produced the rounded l e a f apex. This l a t t e r condition appears to be produced as a response to high humidity and low l i g h t i n t e n s i t y . 4. LEAF LAMINA There are many characters suitable for analysis i n the broad blade of vegetative leaves, excluding the margin and costa. The areolation i s d i r e c t l y responsible for the size and shape of the l e a f . C e l l size i s extremely v a r i a b l e , being proportionately larger i n larger leaves. Plants of the same taxon with various size l e a f c e l l s from d i f f e r e n t Figures 26-33. Comparison of various ecophenic expressions of Philonotis fontana. The f i r s t three leaves i n each figure were drawn from plants as they appeared i n the f i e l d ; (Scale A) the adjacent two leaves were drawn from the same c o l l e c t i o n a f t e r being cultured at the same time i n a uniform environment (Scale B). 26 Scale A Scale B Figure 34. Leaves of Philonotis glauoescens having an acute apex i n the f i e l d (Florida, Zalesj 206 8) developed an obtuse apex i n culture. Figure 35. Leaves of Philonotis glauoescens having an obtuse apex i n the f i e l d ( Florida, Zales} 2041) continue to have an obtuse apex i n culture. These same cultured plants developed leaves with an acute apex when transplanted to the f i e l d , but developed an obtuse apex when returned to the culture chamber. 27 28 environments i n the f i e l d develop the same size c e l l s i n culture. C e l l shape i s consistent and a more useful diagnostic character than c e l l s i z e . Some species have narrow very elongate c e l l s (those with triangular-lanceolate leaves), and others have rectangular, quadrate, to s l i g h t l y i r r e g u l a r c e l l s (those with ovate-lanceolate leaves). These features remain constant i n culture. C e l l wall thickness varies d i r e c t l y i n response to environmental changes. A l l species of Philonotis growing i n wet or densely shaded habitats or both w i l l produce th i n walled lax c e l l s . This lax condition can be produced by growing plants i n water culture. Philonotis laxa and varie t y laxa (taxa based on the lax condition of several species) are ecophenic and should not be recognized taxonomically. The l e a f c e l l p a p i l l o s i t y i s an important family c h a r a c t e r i s t i c and also i s the most stable s p e c i f i c character i n the genus. The p a p i l l a e may be found at the upper, middle, or lower c e l l ends of certain species ( f i g . 36-43), never at both ends on the same surface as implied by some authors and i l l u s t r a t o r s . The leaves must be cleared with lacto-phenol i f an accurate assessment of pa p i l l a e p o s i t i o n i s to be made. Regardless of a l l other v a r i a t i o n i n Philonotis, the p a p i l l a e p o s i t i o n on l e a f c e l l s remains consistent when cultured plants are compared to the same plants i n nature. The size of i n d i v i d u a l p a p i l l a e i s generally smaller i n 29 cultured material due to thinner c e l l walls. Occasionally p a p i l l a e w i l l occur at opposite ends of a c e l l on opposite surfaces of a l e a f . In lo n g i t u d i n a l section the two c e l l s appear to be overriding each other ( f i g . 39). Longifttudinal folds or p l i c a e are evident i n some species. This feature has been employed as an important s p e c i f i c character but, as demonstrated by culture experiments ( f i g . 26-30), i s not always present throughout the same plant. P l i c a t i o n i n the le a f lamina i s a v a l i d character only i n f e r t i l e or very robust plants. C e l l contents have ra r e l y been used for taxonomic purposes. The siz e and number of chloroplasts are variable. Plants i n culture w i l l produce fewer but larger chloroplasts per c e l l than those i n the f i e l d . O i l bodies are occasionally produced but exhibit no c o r r e l a t i o n to the environment, age, or stature of the plants i n culture or i n the f i e l d . Furthermore, the chloroplasts and o i l bodies are lo s t during lacto-phenol treatment. C e l l chemistry may be of diagnostic importance i f proper a n a l y t i c a l methods are employed. A d i s t i n c t i v e a l a r region i s not d i f f e r e n t i a t e d i n Philonotis. 5. LEAF MARGIN Two features of the le a f margin are e a s i l y detectable; they are p a p i l l o s i t y and i n r o l l i n g . P a p i l l o s i t y i s an important and stable taxonomic character. The p a p i l l a e may be at the upper c e l l end only i n some species ( f i g . 42, 43), Figures 36-40. Types of lamina c e l l s found i n the North American species of Philonotis i n surface view and longitudinal section. -36. P. longiseta -37. P. marchica -38. P. capillaris -39. P. fontana -40. P. yezoana Figures 41-43. Types of l e a f margin c e l l s found i n the North American species of Philonotis . -41. P. fontana -42. P. marchica -43. P. (j-Zauceseens 31 or at both ends forming "zwillingsmamillen" of Janzen (1921) ( f i g . 41). These "zwillingsmamillen" or double teeth should not be confused with margins that appear to be doubly toothed due to a r o l l e d margin ( f i g . 43). The presence or absence of double teeth i s apparent i n large f e r t i l e plants but less obvious i n smaller or cultured plants. This s i z e difference i s not important, but t h e i r occurrence i s consistent i n the f i e l d as well as i n culture and of basic taxonomic value. Many small s t e r i l e plants c o l l e c t e d i n nature w i l l show f l a t l e a f margins. These same plants would have produced r o l l e d margins i f they had continued to develop into robust plants. This ecophenic v a r i a t i o n i s demonstrated by the fact that r o l l e d l e a f margins are absent from most species i n culture but present i n the material c o l l e c t e d in situ. ( f i g . 26-32). The presence or absence or degree of i n r o l l i n g of the margin i s too variable to be of much, taxonomic value. 6. COSTA Costa anatomy i n Philonotis i s r e l a t i v e l y consistent. A l l species have a strong costa that reaches nearly to the apex, i s percurrent, or excurrent. In cross section the costa i s composed of one, two, or three guide c e l l s surrounded by several layers of incrassate s t e r e i d s . Cultured plants i n general w i l l have fewer stereids with thinner c e l l walls than the same plants from the f i e l d . The i n t r a s p e c i f i c v a r i a t i o n observed i n costa anatomy i s too great to be employed as an i n t e r s p e c i f i c character. 7. LEAF ARRANGEMENT Phyllotaxis and spacing i s variable throughout Philonotis Plants growing i n a uniform environment show a regular s p i r a l phyllotaxy. Regular geometric phyllotaxis i s rarely observed i n nature, instead the leaves may appear homomallous, somewhat five-ranked, i r r e g u l a r to scattered, or sometimes arranged i n v e r t i c a l rows. The spacing i s consistent i n culture but highly variable i n nature. Very wet or shaded conditions w i l l produce stems with greater distances between the leaves (etiolated p l a n t s ) . Considering t h i s evidence, these modifications are a l l ecophenic and i n v a l i d as taxonomic characters. 8. STEM Stem structure i s r e l a t i v e l y uniform i n t h i s genus. There i s an outer layer of i n f l a t e d c o r t i c a l c e l l s surrounding an area several c e l l s i n thickness composed of very thick walled c e l l s . The center of the stem i s composed of large t h i n walled c e l l s and a central strand of small thin walled c e l l s ( f i g . 4 ) . The stems may be larger with a more prominent central strand i n robust plants when compared to the same plants i n culture. Pigmentation i s more evident i n older and h e a l t h i e r stems than i n young, shaded, aquatic, or cultured plants. 9. RHIZOIDS Rhizoids are too variable to be of taxonomic importance. Their presence i s influenced by moisture. Plants growing i n a dry atmosphere or i n extremely wet to aquatic habitats w i l l 33 have few stem rhi z o i d s . Plants growing i n a humid atmosphere, as i n the cultures or i n some habitats i n nature, w i l l produce an abundant growth of tomentum. The SEM has shown that the fine structure of Philonotis rhizoids i s also var i a b l e . Some parts of the rhizoids may be smooth while others from the same stem may be quite scabrous ( f i g . 44, 45). 10. PROPAGULA Propagula of some form occur on a l l species of Philonotis. They are not consistently present but develop i n response to changing environmental conditions. Abundant propagule production can be induced by allowing plants i n culture to slowly desiccate. In nature as i n the cultures, propagula are important asexual reproductive organs. They occur most frequently on members of the section Philonotula. Those found i n the section Philonotis may f a l l o f f and function as propagules or remain attached and develop into the t y p i c a l whorl of small branches or innovations. Taylor (1958) has described small r e s i d u a l leaves that are present at the point of propagule attachment. These ti n y structures are not unusual i n any plants developing propagules, but are ecophenic and occur too sporadically to be of diagnostic value. D. CONCLUSIONS Philonotis, a common and widespread genus i n North America, can tolerate a great d i v e r s i t y of s o i l and rock types, moisture regimes, and l i g h t i n t e n s i t i e s . As i s true of many Figures 44, 45. Scanning electron micrographs of Philonotis fontana rhizoids produced on the same stem. other waterloving plants, the broad e c o l o g i c a l amplitude of Philonotis i s p a r a l l e l e d by i t s extreme morphological p l a s t i c i t y . Table II l i s t s those characters that are t y p i c a l of a l l species i n the genus, those that are s p e c i f i c for certai n species or groups of species, and those characters that are variable depending on environmental conditions. The nature of stem anatomy, presence of rhiz o i d s , and lanceolate leaves with toothed margins and p a p i l l o s e rectangular c e l l s are important gametophytic c h a r a c t e r i s t i c s that, i n combination, di s t i n g u i s h Philonotis from other genera. The shape of these lanceolate leaves and rectangular c e l l s i n various combinations with the pos i t i o n and number 'of p a p i l l a e are stable characters that are always present and can be used to describe and i d e n t i f y the North American taxa. The size of leaves, c e l l s , and stems; the abundance of r h i z o i d s , propagula, p l i c a e , and r o l l e d margins; or leaf arrangements are a l l variable ecophenic modifications. These ecophenic modifications have, i n the past, been used to recognize taxa with formal taxonomic status. This has caused much confusion and ambiguity i n the understanding of the genus. Table III l i s t s the descriptive names and t h e i r meanings, without author c i t a t i o n s , that have been applied to one or more of the recognized taxa of Philonotis. Monkemeyer (19 27) stated; "Solche Wuch-, Farben-oder Anpassungsformen TABLE II, MORPHOLOGICAL CHARACTERS OF PHILONOTIS ECOPHENIC CONSISTENT THROUGHOUT GENUS CONSISTENT WITHIN SOME SPECIES lax c e l l s p l i c a t e leaves l e a f arrangement r o l l e d or f l a t margin presence of propagulae l e a f s i z e s i z e & color of stems....stem cross section amount of rhizoids presence of rhizoids c e l l s i z e .. ..± rectangular c e l l s elongate rectangular vrs. presence of pa p i l l a e toothed margin lanceolate leaves... quadrate rectangular p o s i t i o n of p a p i l l a e single teeth vrs.double teeth ovate-lanceolate vrs. triangular-lanceolate s o l l t e man der Einfachheit wegen, um den nebensachlichen Formelfram der Benennung und P r i o r i t a t s f r a g e n umgehen zu konnen, je nachdem . . . und . . . ohne Autorzitierung bezeichnen, s i e sind s o f o r t ohne langatmige Beschreibungen verstandlich und das i s t j a die Hauptsache." Buch (1928) also has proposed a s i m i l a r system for designating ecophenic v a r i a t i o n , the modificatio, i n the leafy hepaticae. These proposals are p r a c t i c a l and the use of descriptive adjectives to point out various ecophenic expressions within a taxon should be based on simple e c o l o g i c a l and morphological observations but should avoid the r i g i d i t y of taxonomic nomenclature. Crundwell (19 70) suggests that modificatio be used for variants with no genetic basis because taxa at t h i s l e v e l are often not mutually exclusive. TABLE III. DESCRIPTIVE NAMES REFERRING TO ECOPHENIC MODIFICATIONS M O D I F I C A T I O (descriptive name) Referring to the siz e of tenera mollis t e r r e s t r i s compacta ampliretis gracilescens tenuis laxa A P P E A R A N C E the plant d e l i c a t e , e t i o l a t e d s o f t , f l a g e l l a form small, widely spaced dense tomentose t u f t s splendid, large erect stems slender t h i n , narrow e t i o l a t e d Referring to the leaves f a l c a t a torquata adpressa laxa l a x i f o l i a pumila heterophy11a dimorphophylla b r e v i f o l i a l o n g i f o l i a bent, usually to one side twisted close to the stem i n f l a t e d c e l l s with t h i n walls i n f l a t e d c e l l s with t h i n walls small dimorphic, blunt apex grading i n t o acute dimorphic, acute apex grading i n t o blunt small leaves long leaves Referring to asexual reproductive structures gemmiclada gemmae present propagulicaulis gemmae present Referring to sexuality polygama synoicous 39 VII. SPECIES CONCEPT Now that ample experimental data are available to demonstrate the phenotypic p l a s t i c i t y within the North American species of Philonotis, i t i s possible to exclude formal taxonomic recognition of ecophenic modifications. For the purpose of the present study a species i s considered to be a series of biotypes that can be recognized by a combination of stable morphologically d i s t i n c t characters. A variety i s a closely related, although morphologically and geographically segregated series of biotypes within a species. Modificatio a d j e c t i v a l names may be added to describe variable conditions within a species or variety, but they should be used only for convenience or to c l a r i f y the appearance of various ecophenes or growth forms as such are names without formal taxonomic status. VIII, TAXONOMIC TREATMENT ' A. DESCRIPTION OF THE GENUS IN NORTH AMERICA Genus Philonotis B r i d . Bryol. Univ. 2:15. 1827. Derivation of the name Philonotis i s from the Greek philo - to love and notis = moisture, a l l u d i n g to the habitat of most species. The North American plants are small and slender (about 1 cm long), or robust (to 10 cm), bright green to yellowish, almost prostrate or usually ascending, i r r e g u l a r l y branched or more frequently with whorled f a s t i g i a t e innovations below the gametangial heads, i n loose or usually dense t u f t s forming small patches or extensive sods, the stems clothed i n ± dense reddish tomentum with a prominent central strand and a single layer of v e s i c u l a r t u r g i d c o r t i c a l c e l l s . Small reproductive branches are occasionally present and subtended by minute, ecostate, ovate residual leaves. Leaves arranged i n f i v e to many ± s p i r a l rows, widely spaced to usually crowded, appressed-imbricate to erect spreading, narrowly triangular-lanceolate to broadly ovate-lanceolate, 0.5-3.5 mm long, + p l i c a t e , sometimes f a l c a t e , acuminate and acute or occasionally b l u n t l y ovate. Margins variously serrulate with single or double teeth nearly throughout, i n r o l l e d , or f l a t . Costa subpercurrent to excurrent. Lamina unistratose, except for p e r i g o n i a l leaves, composed of c e l l s variously elongate rectangular to quadrate rectangular, somewhat rounded i n lax plants. P a p i l l a e at upper or lower c e l l ends or middle, on one or both surfaces, 41 r a r e l y smooth. Some species' have dimorphic stem leaves with t y p i c a l leaves on s t e r i l e and female plants and appressed, smaller, and more distant leaves with blunt t i p s on male plants. Dioicous, (monoicous i n P. longiseta), sporangia a r i s i n g terminally becoming l a t e r a l as the stems develop, perigonia gemmiform (bud shaped) or d i s c o i d (rosette shaped), bracts broadly ovate, sheathing at the base, abruptly narrowed with weakly developed costae, paraphyses abundant and f i l i f o r m . Seta s i n g l e , elongate 1-7 cm long, s t r a i g h t , or flexuose. Calyptra small, glabrous, cucullate, fugacious. Capsules ovoid to subglobose, bright green when developing becoming dark rust brown, strongly i n c l i n e d to ho r i z o n t a l , asymmetric, ribbed, or strongly furrowed l o n g i t u d i n a l l y and constricted when empty and dry, short necked with numerous stomata, operculum convex-conic or bl u n t l y apiculate, annulus absent. Peristome double, sixteen teeth i n each c i r c l e , exostomial teeth lanceolate, dark reddish or orange brown, appearing trabeculate with prominent horizontal lamellae occasionally with round t o r i i n the upper segments, segments of the endostome pale yellowish, s t r i a t e p a p i l l o s e , or smooth, with a well developed basal membrane, c i l i a 1-3, well developed, i r r e g u l a r , and occasionally cohering. Spores globose or reniform, densely p a p i l l o s e , brown to orange brown, 20-35y. Hygrophytic mosses growing on various substrates rarely containing much organic material, some species frequenting shaded ditches, c l i f f s , or ravines and others most often i n exposed meadows, lake and r i v e r banks, or springs. Type species: Mnium fontanum (= Philonotis fontana (Hedw.j Brid.) Hedw., Sp. Muse. 195. 1801. 43 B. KEY TO THE NORTH AMERICAN SPECIES OF PHILONOTIS (Leaves must be cleared i n lacto-phenol) 1. Leaf c e l l s p a p i l l o s e at upper or lower ends 2 1. Leaf c e l l s p a p i l l o s e i n the center throughout P. yezoana p. 90 2. Leaf c e l l s p a p i l l o s e at. the upper ends throughout.... 3 2. Leaf c e l l s p a p i l l o s e at the lower ends at least i n lower portion of leaf.. 7 3. Leaf c e l l s elongate (6-20:1) i n + v e r t i c a l rows..4 3. Leaf c e l l s quadrate (1:1) to rectangular (4:1) i n less obvious v e r t i c a l rows 5 4. Leaves long triangular-lanceolate, s t r a i g h t or fa l c a t e and f l a t , c e l l s long and narrow (9-15:1), p a p i l l a e pointed at the extreme upper c e l l ends, monoicous P. longiseta p. 52 4. Leaves tr i a n g u l a r to s l i g h t l y ovate-lanceolate, carinate, c e l l s rectangular (less than 9:1), p a p i l l a e round near the upper c e l l ends, dioicous . P. marchica p. 63 5. P a p i l l a e obvious and numerous, margins r o l l e d and strongly scabrous, costa long excurrent, rare, F l o r i d a only P. sphaerocarpa p. 60 5. Pap i l l a e f a i n t , margins + r o l l e d and toothed, costa variable 6 6. Costa percurrent or ending short of the apex, margins composed of rounded p a p i l l o s e c e l l s appearing doubly serrate, southeastern U.S.A P. glaucescens p. 46 44 6. Costa long excurrent, margins singly serrate, P a c i f i c Coastal P. oapillaris p. 69 7. Margins doubly serrate, (sometimes very faint) P. fontana 8 7. Margins never doubly serrate.P. oapillaris p. 69 8. Plants usually large (4-10 cm), forming loose t u f t s or sods (very slender i n aquatic or shaded p l a n t s ) , ± tomentose , costa variable, h o l a r c t i c 9 8. Plants small (less than 5 cm), forming compact turfs held together by abundant tomentum, costa excurrent, northern or alpine P. fontana var. pumila p. 86 9. Leaves broadly ovate and shortly lanceolate, widely spaced with several p l i c a e on each side of the costa, stem t i p s twisted, western P. fontana var. amerioana P« 82 9. Leaves variable, ovate-lanceolate + f a l c a t e , rarely with shallow p l i c a e , stem t i p s not twisted, cosmopolitan P. fontana var. fontana p. 73 C. EXPLANATION OF SPECIES FIGURES To f a c i l i t a t e reference and comparison of the i l l u s t r a t i o n s , a l l figures have been arranged and l e t t e r e d uniformly. The d e t a i l s of the l e a f cross section, lamina, margin, basal angle, and apex were a l l drawn at the same magnification and reduced i n proportion to the indicated measurement on each plate. A. Leaf B. Median le a f cross section C. C e l l s i n upper portion of lamina D. C e l l s i n median portion of lamina E. C e l l s of l e a f base F. Median marginal l e a f c e l l s G. Apical l e a f c e l l s 46 Philonotis glauoescens (Hornsch.) Broth., Bih. K. Svensk. Vet. Ak. Handl. 21 Afd. 3(3):27. 1895. F i g . 49. Bartramia glauoescens Hornsch. in Mart., F l . B r a s i l . 1(2):40. 1840. TYPE: " i n t r u c i s vetustis silvarum prope Sebastianopolin. B r a z i l . Beyrioh," (Holotype BM seen). Philonotis glauoescens var. brevifolia Flow, in Grout, Moss. F l . No. Amer. 2(3):168. 1935. nom. nud. P. glauoescens f. laxa Flow, in Grout, Moss F l . No. Amer. 2 (3) -.168. 1935. TYPE: "F l o r i d a , Smith," (type reported by Flowers to be at US but not locatable). syn. nov. Bartramia tenella C. Mull., Syn. Muse. Frond. 1:481. 1849. TYPE: "Insula T r i n i t a t i s Antillarum, Cruger," (not seen). Philonotis tenella (C. Mull.) Jaeg., Adumb. 1:541. 1873-4. P. muehlenbergii var. tenella B r i d . , Bryol. Univ. 2:23. 1827. P. sphaerooarpa (Hedw.) Bri d . ssp. tenella (C. Mull.) Jaeg. var. terrestris Dism., B u l l . Soc. Bot. France 10. Mem. 17:14. 1910. TYPE: "F l o r i d a , Beresford, Hulst 6a & 6b," (not seen). P. glauoescens var. terrestris (Dism.) Flow, in Grout, Moss F l . No. Amer. 2(3):168. 1935. syn. nov. P. uncinata var. glauoescens (Hornsch.) Florsch., F l . Suriname 6(1):205. 1964. syn. nov. 47 P. graoillima Angstr. , Oefv. Kgl. Sv. Vet.-TAkad. Forh. 33(4):17. 1876. TYPE: " B r a z i l , Caldas. A. F. Regnell 38," (Holotype S-PA seen). syn. nov. P. unoinata var. graoillima (Angstr.) Florsch., F l . Suriname 6(1):205. 1964. syn. nov. Plants 0.5-2.0 cm, mostly small pale glaucous green to yellowish green forming th i n and scattered patches on s o i l , r a r e l y very tomentose. Leaves 0.4-1.2 mm long (mostly 0.7-8.0) by 0.1-0.3 mm wide, crowded at the top to widely spaced below, erect becoming erect spreading when moist, ovate-oblong or triangular-lanceolate occasionally subfalcate; apex bl u n t l y rounded-obtuse (in young leaves and i n plants of shaded sites) to slenderly acute; costa percurrent or ending below the apex, p a p i l l o s e near the apex on the abaxial surface; margins of one or two rows of i n f l a t e d c e l l s , b l u n t l y p a p i l l o s e at the upper ends only and appearing doubly serrate, r a r e l y f l a t ; lamina of p e l l u c i d , usually t h i n walled oblong-rectangular to rhomboidal c e l l s (2-5:1) becoming quadrate below, rounded p a p i l l a e at upper c e l l ends r e s t r i c t e d to the adaxial surface, usually throughout but occasionally f a i n t or absent. Dioicous, rarely with sporophytes i n North America, p e r i g o n i a l leaves long acuminate forming a bud enclosing the antheridia (gemmiform). Seta 1-2 cm long; capsule 1.5-2.0 mm, sp h e r i c a l ; stomata 50-100 at base of capsule; outer peristome teeth as for the 48 genus, occasionally producing spherical i n t e r l a m e l l a r thickenings, inner peristome evenly p a p i l l o s e ; spores 22-26y b l u n t l y rounded p a p i l l o s e . CHROMOSOME NUMBER: n = 6. J. Snider has generously permitted i n c l u s i o n of the report of t h i s count he made at Duke University from a culture of P. glaucescens ( F l o r i d a , Hernando Co., Bro o k s v i l l e , Zales 2041 UBC) ( f i g . 51). HABITAT: On basic mineral substrata (limestone, sandstone, sandy s o i l , mud, or rarely charred wood), i n ditches or sinkholes, along r i v e r or creek banks, and at the margins of springs, w a t e r f a l l s , or dams, predominantly i n shaded locations. DISTRIBUTION: Common i n the northern two thirds of Fl o r i d a and rare i n Texas, Louisiana, and Georgia. Reported from the West Indies, Mexico, Central and South America (Map f i g . 6 4 ) . Philonotis glaucescens and P. gracillima were distinguished s o l e l y on the basis of le a f apex shape and costa length. Most of the c o l l e c t i o n s examined, including the type specimens of both species, show both rounded-obtuse leaves with the costa ending just below the apex and acuminate leaves with a percurrent costa. The separation of these two taxa was usually a subjective judgement dependent on which kind of le a f was more abundant. Culture experiments demonstrate that these two characters are ecophenic ( f i g . 34, 35) . Occasionally P. glaucescens has been confused with Figures 4 6-48. Comparison of leaf shape and leaf margin -46. Philonotis glauoescens -47. Philonotis marchica -48. Philonotis sphaerooarpa 50 P. marchica. Very young and lax P. marchica may have a blunt apex with f a i n t p a p i l l a e on the lamina but the triangular-lanceolate shape and f l a t l e a f margin of oblong pointed teeth w i l l d i s t i n g u i s h i t from the lanceolate shape and usually r o l l e d l e a f margin composed of short, blunt, and somewhat i n f l a t e d teeth of P. glaucescens ( f i g . 46, 4 7 ) . Florschutz (1964) has treated P. glaucescens and P. graoillima as v a r i e t i e s of P. uncinata with the reservation that even v a r i e t a l status may be too high. Because P. sphaerocarpa does not occur i n the area of Florschutz' monograph he missed the opportunity of synonymizing i t with P. uncinata. Philonotis sphaerocarpa however has p r i o r i t y . He i s correct i n combining P. graoillima and P. glauces cens but overlooked several d i s t i n c t i v e features separating them from P. sphaerocarpa (uncinata). Philonotis glaucescens i s a small pale plant rarely exceeding 2.0 cm with ovate-oblong or triangular-lanceolate leaves of usually thin walled c e l l s with f a i n t p a p i l l a e and b l u n t l y toothed margins. Philonotis sphaerocarpa i s generally robust, yellowish green, and 2.0-5.0 cm high with concave oblong-lanceolate leaves of thick walled c e l l s with prominent p a p i l l a e and a scabrous appearing margin due to the large p a p i l l a e on the revolute laminar margin ( f i g . 46, 48). Figure 4 9 . Philonotis glaucescens 52 Philonotis longiseta (Rich, 'in Michx.) B r i t t . , The Bryologist 14:44. 1911. F i g . 54. Bartramia longiseta Rich, in Michx., F l . Bor. Amer. 2:301. 1803. TYPE: " i n Carolina ex D. Bosc et i n V i r g i n i a ex D. Beauvois," (Isotype NY seen). Philonotis longiseta var. propagulaecaulis Flow., The Bryologist 38:8. 19 35. TYPE: "wet s o i l , Louisiana, Lafayette, Leon 1353," (Holotype US seen). syn. nov. P. longiseta f. propagulioaulis Flow, in Grout, Moss F l . No. Amer. 2(3):172. 1935. nom. nud. P. longiseta var. polygama Flow, in Grout, Moss F l . No. Amer. 2(3):171. 1935. TYPE: "near C a r e y v i l l e , Campbell Co., Tennessee, on moist roadside bank, Sharp 34154," (Holotype UT seen). syn. nov. Bartramia radicalis P. Beauv., Prodr. Aetheog. 44. 1805. Philonotis radicalis (P. Beauv.) B r i d . , Bryol. Univ. 2:17. 1827. Bartramia radicalis var. porteri Aust., B u l l . Torrey Bot. Club 6 (36) :190. 1877. TYPE: "on wet rocks and banks, western Pennsylvania, Garter', Rockdale (Lehigh Valley) Wolle 1874; Oneida, New York, Warne 1873; Ohio, Beardslee & Biddlecome," (Lectotype Wolle 1874 F seen). Philonotis longiseta var. porteri (Aust.) Flow, in Grout, Moss F l . No. Amer. 2(3):171. 1935. syn. nov. Plants 0.5-3.0 cm high (mostly 2.0), pale yellowish green to green, forming loose to dense, usually small rounded t u f t s , branching e s p e c i a l l y below the sex organs, 53 usually rust brown tomentose, often producing deciduous propaguliferous shoots. Leaves 1.0-2.0 mm long, by 0.1-0.3 mm wide, erect and occasionally secund when dry becoming spreading when moist, l i n e a r triangular-lanceolate to lanceolate, slenderly acuminate with an excurrent costa; c e l l s long l i n e a r (9-15:1) in. v e r t i c a l rows becoming gradually enlarged below, not i n f l a t e d , p a p i l l o s e at the extreme upper ends of a l l c e l l s on the adaxial surface only; margins plane or narrowly revolute + the en t i r e length usually on both sides,, strongly serrate with blunt teeth at upper c e l l ends only. Autoicous or synoicous, perigonia gemmiform (composed of erect leaves enclosing the gametangia and forming a "bud"). Seta 1.5-3.5 cm long, capsule rounded, stomata 67-146 (average 100) per capsule, r e s t r i c t e d to the lower one quarter, peristome as for the genus, spores 22-33y, reniform, p a p i l l o s e . CHROMOSOME NUMBER: n = 12. Heitz (1942) and Lowry (194 8) reported f o r the genus Mnium, that those species which have the basic chromosome number are generally dioicous and those species which are polypoids are monoicous. This also seems to be the case i n Philonotis. Ochi (1963) has shown that the dioicous and monoicous races of P. socia i n Japan are the same and Yano (1955) has described n = 6 and n = 12 populations of P. socia, but i t i s not clear from the l i t e r a t u r e i f the dioicous populations are n = 6 and the monoicous populations are n = 12. This generalization i s supported by the monoicous P. longiseta. J . Snider, using the f a c i l i t i e s at Duke University, has established a chromosome number of n = 12 f o r a polygamous specimen of P. longiseta (Florida, Leon Co., Tallahassee, Zales 1941 UBC) ( f i g . 50). A l l of the species of Philonotis that have been counted except for P. socia and P. longiseta are dioicous and n = 6. I r o n i c a l l y there i s no n = 6 dioicous taxon that corresponds morphologically to P. longiseta, at le a s t not i n North America. HABITAT: On sandstone, limestone, or phosphate rock, sandy or clay s o i l low i n organic material, i n ditches, ravines, or on r i v e r banks and b l u f f s , around springs, or along roadsides. Usually at low elevations i n shaded places that are wet for some part of the year. DISTRIBUTION: Predominantly southeastern, common i n the Ozarks and southern Appalachian mountains extending north along the r i v e r s into Iowa, west into central Texas, south to central F l o r i d a , and northeast to Pennsylvania. Reported from Guatemala, Martinique, Mexico, and Puerto Rico (Map f i g . 65). Philonotis longiseta i s a c l e a r l y demarcated species, geographically, sexually, c y t o l o g i c a l l y , and morphologically. In North America P. longiseta occurs east of the 100th meridian and mostly south of the 40th p a r a l l e l . I t i s the only North American species of Philonotis that i s monoicous or has a haploid chromosome number of 12, and i t has long lanceolate-triangular leaves with revolute margins and Figures 50, 51. M i t o t i c chromosome configurations of Philonotis -50. P. longiseta -51. P. glauoes cens 55 56 p a p i l l a e at the extreme upper ends of most c e l l s . The p e r i g o n i a l leaves are variable and of l i t t l e taxonomic value. They consist of broad, lax, almost ecostate inner leaves that give the perigonium an almost d i s c o i d appearance to leaves that are s i m i l a r to enlarged comal leaves. Philonotis longiseta has occasionally been confused with P. mavchioa. I f the leaves are cleared with lacto-phenol i t i s obvious that the former has longer narrower c e l l s with p a p i l l a e at the extreme upper c e l l ends, and the l a t t e r has p a p i l l a e on or near the upper c e l l ends. Philonotis mavehiaa also has a shorter broader l e a f . Extremely young and lax or e t i o l a t e d plants may have shorter and broader leaves with s l i g h t l y rounded c e l l s , plane margins, and a percurrent costa. These can be recognized as P. longiseta by the p o s i t i o n of the p a p i l l a e on the l e a f c e l l s ( f i g . 36, 37, 52, 53). Philonotis longiseta var. porteri was described from several c o l l e c t i o n s possessing erect, s l i g h t l y tapering leaves, shorter stems and seta, and a more erect capsule. A l l but one of the specimens c i t e d i n Austin's description (1877) of Bartramia radicalis var. porteri have been seen: western Pennsylvania, Garber (not seen); Rockdale, Lehigh Valley, Vlolle 1874 (Lectotype F) i s a lax expression of the species; Oneida, New York, Warne 1873 (NY) i s P. longiseta, 1874 (NY) i s a mixture of P. longiseta and P. marchica', Ohio, Beardslee (NY) i s P. longiseta, Biddlecome 1877 (US) i s P. marchica. Figures 52, 53. -52. -53. Comparison of l e a f shape and median le a f c e l l shape Philonotis marchica Philonotis longiseta Besides the o r i g i n a l description, several pages of Austin's notes are housed i n the New York Botanical Garden Herbarium which compare B. radicalis var. porteri with B. striata and P. marchica but give no i n d i c a t i o n of i t s difference from B. radicalis . The minor si z e differences seen i n the syntypes do not support the recognition of th i s variety as being d i s t i n c t from the species. 59 Figure 54. Philonotis longiseta 60 Philonotis sphaerocarpa (Hedw.) Brid., Bryol. Univ. 2:25. 1827. F i g . 55. Mnium sphaericarpon Hedw., Sp. Muse. 197. 1801. TYPE: "In Jamaica, Swartz," (Isotype NY seen). Philonotis sphaericarpa var. terrestris Flow, in Grout, Moss F l . No. Amer. 2(3):168. 1935. nom. nud. Bartramia uncinata Schwaegr., Suppl. Sp. Muse. 1(2):60. 1816. TYPE: "In monte sulphurifero Guadalupae et in Martinica lectum et curiose observatum dedit, Richard," (Holotype Guadalupae PC seen). Philonotis uncinata (Schwaegr.) Brid., Bryol. Univ.. 2:22. 1827. syn. nov. Plants 2.0-5.0 cm, r e l a t i v e l y robust, yellowish green erect t u f t s or dense mats, abundantly rust brown tomentose below, often branched below the sex organs. Leaves 1.0-2.0 mm long, usually crowded, erect or erect-spreading, s t r a i g h t or subfalcate e s p e c i a l l y at the stem t i p s , lanceolate to oblong-lanceolate often concave at the base, acuminate with the costa short excurrent to strongly spinulose-excurrent, margins usually strongly revolute appearing doubly or more serrate nearly to the base, upper c e l l s l i n e a r or oblong-linear (3-7:1) becoming short rectangular below, densely scabrous with p a p i l l a e at upper c e l l ends on the adaxial surface. Dioicous, perigonia small and bud shaped (gemmiform). Seta 1.5-3.0 cm long, capsule spherical, 1.5-2.5 cm, stomata 90-150 r e s t r i c t e d to the base, peristome as for the 61 genus, spores 20-26u p a p i l l o s e . CHROMOSOME NUMBER: Not reported. HABITAT: On damp s o i l and rock banks and c l i f f s at various elevations i n the American t r o p i c s . DISTRIBUTION: One st a t i o n i n North America (Sanford, F l o r i d a ) , West Indies, Mexico, Central and South America (Map f i g . 66). Examination of the type material of P. sphaerooarpa and P. uncinata and many authentic specimens from the West Indies, Mexico, Central and South America indicate that the range of v a r i a b i l i t y of these taxa overlap to such a great extent that they are impossible to separate. The only major difference between the type specimens i s the amount of curvature i n the leaves which i n the genus Philonotis i s a notoriously variable character. The single waif specimen from North America was col l e c t e d by S. Rapp i n 1905 i n a hummock near Sanford, F l o r i d a . A l l other specimens i d e n t i f i e d as P. sphaerooarpa have been assigned to P. longiseta^ glauoescenssor marchica. For the characters separating P. sphaerooarpa from P. glauoescens, see the discussion under P. glauoescens. 63 Philonotis marchica (Hedw.) Bri d . , Bryol. Univ. 2:23 & 735. 1827. F i g . 58. Mnium marchicum Hedw., Sp. Muse. 196. 1801. Philonotis glabriusoula Kindb. in Mac. et Kindb., Cat. Can. PI. 6:107. 1892. pro. parte. TYPE: "by springs i n a gully, Canaan Forks, Queen's Co., N. B., January 2nd, 1890. J. Moser," (Syntype CAN seen, S-PA seen). P. subcapillaris Kindb., Eur. No. Amer. Bryin. 326. 189 7. pro. parte. TYPE: "Switzerl. near Lugano," (not seen). Bartramia muehlenbergii Schwaegr., Suppl. Sp. Muse. 1(2):58. 1816. TYPE: " i n Pennsylvania, Muhlenberg," (not seen). Philonotis muehlenbergii (Schwaegr.) Br i d . , Bryol. Univ. 2:22. 1827. Plants 0.5-6.0 cm high, mostly small and yellowish t u f t s i n dry habitats, becoming robust and brighter green forming erect mats i n moist areas and high elevations. Stems brown tomentose below and occasionally clothed with abundant propagula. Leaves 0.9-2.2 mm long by 0.3-0.5 mm wide, erect spreading, more divergent when moist, occasionally arranged i n f i v e s p i r a l rows i n robust plants, triangular-lanceolate mostly s t r a i g h t , carinate; margins plane or only narrowly recurved, serrate nearly to the base with p a p i l l a e at the upper ends only; costa narrow, shortly excurrent to c l e a r l y 64 excurrent; upper c e l l s linear-oblong (5:1), 4-8u wide, lower c e l l s oblong (3-4:1), 6-15u wide, p a p i l l a e obvious at upper ends of c e l l s throughout, on the adaxial or dorsal surface only. Dioicous, perigonia d i s c o i d (forming a roset t e ) . Seta 2-4 cm long; capsule spherical, 1.0-2.5 mm, stomata two or ra r e l y four c e l l e d , 90-200 mostly 130 or more per capsule, r e s t r i c t e d to the base; outer peristome as for the genus, inner peristome p a p i l l o s e i n l i n e s , c i l i a v a riable; spores 20-30y, p a p i l l o s e . CHROMOSOME NUMBER: Not reported. HABITAT: On rocks and sandy s o i l i n wet places, usually i n slowly running water and forming small to large mats depending on the moisture, at various elevations. Reported on calcareous s c h i s t , limestone, sandstone, t r a v e r t i n e , bogs (?), and roadside ditches. DISTRIBUTION: Rare i n the Rocky Mountains from southern B r i t i s h Columbia and Alberta south to Mexico, absent from the i n t e r i o r plains but reappearing i n the central highlands of Missouri and adjacent states, the mountains of eastern North America from southern Newfoundland to Georgia, and i n scattered l o c a l i t i e s around the Great Lakes. Reported from central and southern Europe and the western Mediterranean (Map f i g . 67). The e x i s t i n g l i t e r a t u r e implies confusion between P. marchica and P. capillaris. However, the habit of th i s species i s usually much more erect and r i g i d than P. capillaris. 65 The stem leaves are closer and more erect and have a triangul a r base compared to the ovate base of P. oapillaris. The l e a f c e l l s of P. marchica are longer, narrower, and d i s t i n c t l y mammillose throughout ( f i g . 56, 57). Besides obvious morphological differences the range of P. marchica and P. oapillaris are exclusive except possibly i n central Idaho and southern B r i t i s h Columbia. Philonotis muehlenbergii (Schwaegr.) Brid. was o r i g i n a l l y described (Schwaegrichen 1816) from a single specimen c o l l e c t e d from Pennsylvania by Muhlenberg. This specimen appears to no longer be i n existence. The species has per s i s t e d i n the l i t e r a t u r e to the present time i n spite of Dismier's review (1910), and the subsequent t r a n s l a t i o n and pub l i c a t i o n of the same paper i n The Bryologist by Br i t t o n (1911), which demonstrated the lack of d i s t i n c t characters separating i t from P. marchica. Flowers (e. a.) has synonymized P. muehlenbergii and P. marchica for the state of Utah; "I have found that those with triang u l a r to oblong-lanceolate leaves, broadest at the i n s e r t i o n , ultimately have shown some of the lower c e l l s to be pa p i l l o s e at the upper ends and referrable to P. marchica." Recent taxonomic works have r e l i e d on the absence or presence of only rudimentary c i l i a to separate P. muehlenbergii from P. marchica. Considering the v a r i a b i l i t y observed i n a large series of specimens throughout t h e i r geographic range and the ease with which the f r a g i l e c i l i a are broken and l o s t , there i s no reason to maintain 66 P. muehlenbevgii as a separate taxon. For a comparison of P. marchica and P. glaucescens see the discussion under P. glaucescens. Figures 56, 57. -56 . -57. Comparison of leaf shape, apex, and median l e a f c e l l s Philonotis marchica Philonotis capillaris 69 Philonotis capillaris Lindb. ex Hartm., Skand. F l . ed. II 10:40. 1871. TYPE: "R. Spruce, Lassaron, Pyreneerna," (Isotype M7ANCH seen, S-PA seen). F i g . 59. Philonotis capillaris Lindb., Hedwigia 6:40. 1867. nom. nud. P. fontana (Hedw.) B r i d . var. capillaris (Lindb.) Lindb., c f . Dism. Soc. Nat. S c i . Nat. Math. Cherbourg Mem. 36:392. 1907. P. macounii Lesq. et James, Man. Mosses No. Amer. 208. 1884. TYPE: "Vancouver Island, Macoun, Can. Musci 152," (Lectotype US seen, Isotype H seen). P. vancouveriensis Kindb., Eur. No. Amer. Bryin. 2:326. 1897. pro. parte. TYPE: "Vancouver Island and Alaska, J. M. Macoun," (Syntype S seen, Isotype CAN seen). P. arnellii Husn., Muse. G a l l . 268. 1890. Plants slender, 1-3 cm high, i n loose t u f t s or scattered, often procumbent, tomentose below. Leaves 0.7-1.5 mm long by 0.3-0.5 mm wide, erect when dry, erect spreading when moist, lanceolate from an ovate base, acuminate, without p l i c a e ; margins plane or narrowly recurved above the ovate base, d i s t i n c t l y serrate nearly to base with single teeth at upper ends of c e l l s only, basal border of f i v e to ten or more quadrate c e l l s ; costa excurrent to long excurrent, upper c e l l s (2-3:1) 6 - l l y wide, lower c e l l s (2-1) 8-14y wide; p a p i l l a e obscure i n lower h a l f of l e a f on the adaxial surface, rarely at lower c e l l ends or 70 overlapping on opposite surfaces, c e l l walls thick, the lumen rounded. Dioicous, often c o l l e c t e d with sporophytes, perigonia d i s c o i d (forming a rosette). Seta 2-3 cm long, capsule rounded, furrowed when dry; stomata 40-115 per capsule, r e s t r i c t e d to the base, mostly less than 100; outer peristome as for the genus, inner peristome p a p i l l o s e i n l i n e s , c i l i a v a riable; spores 20-26u, pa p i l l o s e . CHROMOSOME NUMBER: Not reported. HABITAT: On wet sandy s o i l , humus, or various t h i n s o i l s over rock on slopes, c l i f f ledges, or ditches. P r e f e r r i n g wet shaded places mixed with small vascular plants and other bryophytes or both. DISTRIBUTION: The P a c i f i c Coast from Alaska to central C a l i f o r n i a and east to northern Idaho and southeastern B r i t i s h Columbia. Reported from Europe and Asia Minor (Map f i g . 68) . The type c o l l e c t i o n i s s t e r i l e and immature material c o l l e c t e d by R. Spruce from the Pyrenees. However, i t does demonstrate the s p e c i f i c characters. The margins of P. oapillaris are composed of single teeth, a l l projecting at the upper c e l l ends as compared to the P. fontana complex where the margins are doubly toothed by c e l l s projecting at both ends. Philonotis oapillaris has been reported from several stations i n central and eastern North America, however, these have a l l ultimately shown some c e l l s with two projections, and thus cannot be included i n t h i s species. The p a p i l l a e of the broadest part of the lamina are obscure and must be cleared with lacto-phenol to be observed at a l l . P a p i l l a e are at the upper ends of the c e l l s but occasionally may be i n the lower ends as i n the type material. The small si z e of f e r t i l e plants, long excurrent costa f l a t almost smooth lamina, ovate l e a f base with single toothed margins, and P a c i f i c Coast d i s t r i b u t i o n i n North America make thi s species e a s i l y recognizable. The European P. arnelli may be conspecific with P. oapillaris, as stated by Dismier (1907b, 1910), Brotherus (1923), and Monkemeyer (1927). Certainly a l l North American specimens labeled P. arnellii are P. oapillaris. Philonotis oapillaris occasionally possesses propaguliferous shoots that would normally develop into branches (innovations t y p i c a l of the genus) i f the plant continued to grow. Propaguliferous shoots in. P. oapillaris have also been reported by F i e l d (1965) and Dismier (190 7b) 72 Figure 59. Philonotis capillaris 73 Philonotis fontana (Hedw.) B r i d . var. fontana, Bryol. Univ. 2:18. 1827. F i g . 60. Mnium fontanum Hedw., Sp. Muse. 195. 1801. Bartramia fontana (Hedw.) Turn., Muse. Hib. 107. 1804. Philonotis adpressa Ferg. in Hunt, Mem. L i t . P h i l . Soc. Manchester ser. 3(5):102. 1872. P. fontana var. adpressa (Ferg.) Limpr., Krypt. S i l e s . 1:116. 1876. syn. not). P. fontana ssp. adpressa (Ferg.) Dix., Stud. Handb. B r i t . Moss. 295. 1896. P. aoutiflora Kindb. in R o l l , Hedwigia 35:67. 1896. TYPE: " V i c t o r i a , Vancouver Island, J. Roll 52," (Lectotype S-PA seen, Isolectotype MANCH seen). P. acutifolia Kindb. ex Dism., B u l l . Soc. Bot. France 10, Mem. 17:27. 1910. nom. inval. err. pro. P. aoutiflora Kindb. P. alpioola var. tenuis Kindb. in Moll., The Bryologist 27:75. 1924. nom. nud. P. braohychypus Kindb. in Moll., The Bryologist 27:75. 1924. nom. nud. P. tomentella Mol. f. brachycarpa (Kindb.) Moll., The Bryologist 27:75. 1924. nom. nud. P. fontana var. brachyphylla Kindb. in Mac. et Kindb., Cat. Can. PI. 6:107. 1892. TYPE: "P. E. I., Barkley Pt. i n bogs, 28 June 1888, Macoun," (Isotype CAN seen). 74 P. glabriuscula Kindb. in Mac. et Kindb., Cat. Can. PI. 6:107. 1892. pro. parte. TYPE: "by springs i n a gully, Queen's Co., N. B., Jan. 2, 1890, J. Moser," (Syntype C/AN seen, MANCH seen, WIS seen). P. microcarpa Kindb., Rev. Bryol. 32:37. 1905. TYPE: "eastern slope of Rocky Mts., bogs along Pipestone Cr. a l t a 6,000 f t . , 8 July 1904, 75," (Lectotype S-PA seen). P. fontana var. mi or obi asta C. M. et Kindb. in Mac. et Kindb., Cat. Can. PI. 6:107. 1892. TYPE: "Selkirk Mt. Roger's Pass, along R. R., 30 July 1890, Macoun," (Lectotype S-PA seen, Isolectotype CAN seen). P. fontana var. microthamnia Kindb., Ottawa Nat. 23:183. 1910. TYPE: "Yukon, Hunker Creek, peat bog, July 1902, 152," (Isotype CAN seen). P. fontana var. columbiae Kindb. in Mac. et Kindb., Cat. Can. P i . 6:107. 1892. (published as Columbia) TYPE: "on wet rocks below the I l l i c i l l e w a e t Canyon, Revelstoke, B. C., 18 May 1890, Macoun," (Holotype S-PA seen). P. fontana var. serrata Kindb. in Mac. et Kindb., Cat. Can. PI. 6:107. 1892. TYPE: "Selkirk Mountains, swamps i n Roger's Pass, 24 August 1885, Macoun', Ounalaska & Atkah Islands, Behring Sea, 1891, Macoun," (Lectotype Roger's Pass CAN seen, Isosyntype Ounalaska & Atkah Islands CAN seen). 75 P. fontana var. heterophylla Card, et Ther., Univ. C a l i f . Publ. Bot. 2:300. 1906. TYPE: "Unalaska, Setchell 2 & 30," (Lectotype 2 UC seen, Syntype 30 UC seen), syn. nov. P. vanoouveriensis Kindb., Eur. No. Amer. Bryin. 2:326. 1897. pro. parte. TYPE: "Vancouver Island & Alaska, Macoun," (Syntype S-PA seen, Isosyhtype CAN seen). P. tenella C. Mull. var. ooloradensis Ren. et Card., Bot. Gaz. 22:51. 1896. orth. err. P. venella. TYPE: "Colorado, Boulder Co., Springdale, M. Eolzinger 1892," (Lectotype NY seen, Isolectotype CAN seen, WIS seen). P. fallax Dism., B u l l . Soc. Bot. France 10. Mem. 17:35. 1910. TYPE: "Canada, B. C., Sicamous, Finlayson 1583 . . . Idaho, Lake Pend d ' O r e i l l e , Leiberg 49," (Lectotype Finlayson NY seen, Syntype Leiberg CAN seen, NY seen). P. fontana f. dimorphophylla Flow, in Grout, Moss F l . No. Amer. 2(3):175. 1935. TYPE: "Alaska," (not seen). P. fontana var. seriata B r e i d l . f. oooidentalis Flow. in Grout, Moss. F l . No. Amer. 2(3):176. 1935. nom. nud. P. calcarea (B. S. G.) Schimp. f. oooidentalis Flow, in Grout, Moss F l . No. Amer. 2(3):179. 1935. TYPE: "Texas, Harvard.s. n.," Not i n US as reported by Flowers, (Isotype SMU seen). 76 P. fontana var. flaoata B r i d . , Bryol. Univ. 2:21. 1827. P. caespitosa Jur., Verh. Zool.-Bot. Ges. Wien 11:235. 1862. syn. nov. P. fontana var. caespitosa (Jur.) Limpr., Krypt. F l . Schlesien 1:116. 1876. P. caespitosa var. compacta Dism., B u l l . Soc. Bot. France 10. Mem. 17:28. 1910. TYPE: "Conn., Ledyard & Hamdem, Nichols," (not seen). syn. nov. P. caespitosa var. compacta Jaeg., Ber. S. G a l l . Naturw. Ges. 1873-74:86. 1875. (Ad. 1:548). nom. nud. P. caespitosa var. heterophylla Dism., B u l l . Soc. Bot. France 10. Mem. 17:28. 1910. No specimen c i t e d . P. caespitosa var. adpressa Dism., Rev. Bryol. 34:68. 1907. syn. nov. P. caespitosa var. ampliretis (Dix.) Crum et al., The Bryologist 68:434. 1965. syn. nov. P. fontana var. ampliretis Dix., Jor. Bot. 40:73. 1902. P. caespitosa var. laxa Loeske et Warnst. in Loeske, Hedwigia 45:104. 1906. P. laxa Warnst., Krypt. F l . Mark Brendenb. 2:619. 1895. horn, illeg. (P. laxa Limpr., Laubm. Deutschl. 2:563. 1893. Name not North American = P. marchica). P. fontana var. laxa Flow, in Grout, Moss F l . No. Amer. 2(3):178. 1935. horn, illeg. (P. fontana var. laxa Vent. Rev. Bryol. 9:45. 1882). P. fontana var. laxa f. tenuis Flow, in Grout, Moss F l . No. Amer. 2(3):178. 1935. TYPE: "colle c t e d i n New York, Pennsylvania and Colorado," (Lectotype Holz. Musci Aero. Bor.-Amer. 661 NY seen, Isolectotype CAN seen). P. fontana f. flowersii Crum, The Bryologist 72:244. 1969 . syn. nov. P. fontana var. aristinervis (Monk.) Diener, Cas. Nar. Mus. Praha 104:122. 1930. P. fontana f. aristinervis Monk., Laubm. Eur. 584. 121c. 1927. P. amerioana var. graoilesoens Dism., B u l l . Bot. Soc. France 10. Mem. 17:23. 1910. TYPE: "Canada, Gaspe Co., Mt. Albert, P. Q., J. F. Collins 4172," (Holotype NY seen). syn. nov. Plants extremely variable, 2.0-10.0 cm high, (rarely larger) usually robust erect ± i r r e g u l a r l y branched, often producing whorls of short branches below the sex organs, bright yellow green to green, forming small scattered tufts to extensive dense sods matted with rust brown tomentum. Leaves erect, appressed to erect-spreading ± falcat e and secund, 0.7-2.5 mm long, by 0.5-1.5 mm wide, mostly ovate-lanceolate with a broad base gradually or abruptly tapering to a ± slender acuminate apex (occasionally blunt), costa stout, percurrent to excurrent, margins revolute or plane, serrate with some c e l l s , p r o j ecting at both ends (doubly serrate) becoming plane at the base and singly serrate near 78 the apex, lamina usually concave with one or several shallow p l i c a e on ei t h e r side of the costa, usually plane, upper leaf c e l l s l i n e a r or oblong-linear (4-10:1) papi l l o s e at the upper c e l l ends on the abaxial surface, lower c e l l s becoming shorter and broader and papil l o s e at the lower c e l l ends on the abaxial surface. Male stems usually rust brown with distant, broad, and appressed leaves with blunt apexes and lax c e l l s with f a i n t p a p i l l a e . Dioicous, perigonia d i s c o i d (rosette shaped) bracts large, lax, and highly v a r i a b l e . Seta 2.0-6.0 cm long, capsule rounded, stomata 110-275 r e s t r i c t e d to lower quarter, peristome as for the genus, spores 20-30u, p a p i l l o s e . CHROMOSOME NUMBER: n = 6 Anderson & Crum (1958), Vaarama (1953, 1956), Smith & Newton (1966). HABITAT: Philonotis fontana occurs i n almost any wet habitat, but i s most abundant and luxuriant i n seepage water i n exposed locations such as ditches, sloping banks of r i v e r s or highways, and around springs or small w a t e r f a l l s , also wet meadows and lake shores, and growing over mineral substrates such as mud, sand, gravel, or rocks of various composition. Plants growing i n shaded, dry, or completely aquatic s i t e s show ecophenic v a r i a t i o n . DISTRIBUTION: Widespread throughout most of North America e s p e c i a l l y i n regions of variable topography, generally absent from the northern tundras, central p r a i r i e s , and southern portions of the states bordering on the Gulf of Mexico. 79 Widespread i n Europe, Asia, northern A f r i c a , and extending i n t o Mexico (Holarctic) (Map f i g . 69). Philonotis fontana has a phenotype that i s e a s i l y modified by various environmental conditions. So many of these modified plants have been described and named that p o s i t i v e i d e n t i f i c a t i o n has become d i f f i c u l t and confusing. Experimental cultures have shown that l e a f shape, s i z e , color, arrangement on the stem, thickness of c e l l walls, and presence or absence of tomentum are highly variable ecophenic c h a r a c t e r i s t i c s , and thus useless as diagnostic characters. The names P. adpressa and var. adpressa of several species have been used for plants with broad and short leaves with blunt apexes, c l o s e l y appressed to the stem. This i s a common condition of male or semiaquatic plants growing i n f u l l exposure to the sun. The names tomentella and borealis used at the s p e c i f i c or v a r i e t a l l e v e l (sensu European workers) have been applied to plants that have narrow elongate leaves with slender excurrent costae. According to the habitat data on the specimens t h i s seems to be a response to low temperature and high elevation. These plants often have leaves swept to one side which have been included under descriptions of var. falcata. Philonotis caespitosa and several lesser v a r i e t i e s of P. caespitosa have been described from small plants that usually have f l a t leaves with plane margins. These plants are almost always s t e r i l e and represent small plants of P. fontana. The epithet laxa has been applied by several authors to several species and v a r i e t i e s of Philonotis. The nomenclature i s confused because of d i f f e r e n t i n t e rpretations. They a l l r e f e r to plants growing i n aquatic or semiaquatic habitats that are heavily shaded causing the plants to become e t i o l a t e d , pale i n color, have i n f l a t e d c e l l s with t h i n c e l l walls, and lack tomentum. N . C. Kindberg (1905, 1910) has described several specie and v a r i e t i e s , mostly from western Canada, that are only minor variations of P. fontana. These names are represented by r e l a t i v e l y few specimens throughout national herbaria. Philonotis fontana i s re a d i l y separated from other species of Philonotis by the large size of the plants, the presence of obvious p a p i l l a e at the lower ends of the le a f c e l l s , and the doubly serrate margins. The d i s t i n c t i v e features of v a r i e t i e s americana and pumila are discussed under the respective taxa. 82 Philonotis fontana var. amerioana (Dism.) Flow, ex Crum, The Bryologist 72:244. 1969. F i g . 61. P. amerioana Dism., B u l l . Soc. Bot. France 10. Mem. 17:35. 1910. TYPE: "high up on the Selkirk Mts., near Armstrong, B. C., Wilson 19 July 1904 as P. fontana," Holzinger, Musci Aero. Bor.-Amer. 70b, (Lectotype NY seen, Isolectotype CAN seen). P. seriata ssp. amerioana Dism., B u l l . Soc. Bot. France 10. Mem. 17:22. 1910. P. torquata Geh., Rev. Bryol. 23:61. 1896. P. amerioana var. torquata (Ren. et Geh.) Dism., B u l l . Soc. Bot. France 10. Mem. 17:23. 1910. P. macounii var. torquata Ren. et Geh., Rev. Bryol. 23:61. 1896. TYPE: "Washington, Olympia, Blacker 40," (not seen). P. amerioana f. laxa Flow, in Grout, Moss F l . No. Amer. 2(3):178. 1935. TYPE: "Selkirk, Canada," (not seen). P. amerioana f. dimorphophylla Flow, in Grout, Moss F l . No. Amer. 2(3):178. 1935. TYPE: "Alaska, under snow banks i n cold water," (not seen). Largest of the North American species of Philonotis, stem 5-10 cm or more long, erect, simple, or with whorled branches, interwoven with brown tomentum, forming deep yellowish green t u f t s or mats. Leaves usually distant, c h a r a c t e r i s t i c a l l y diverging from the stem at a wide angle with the upper portion curving upward, often twisted forming a s p i r a l l i n g stem t i p , 83 broadly ovate, concave, s t r a i g h t , or falcate, shortly acuminate with a percurrent or excurrent costa; lamina p l i c a t e with rounded rectangular c e l l s p a p i l l o s e at the lower end becoming longer near the l e a f apex; margins usually recurved and strongly doubly serrate. Dioicous, perigonia large, d i s c o i d (rosette shaped). Seta 3.0-6.0 cm long; capsule rounded, furrowed when dry; stomata 130-2 30 r e s t r i c t e d to the base of capsule; peristome and spores as for the species. CHROMOSOME NUMBER: Not reported. HABITAT: Occurring i n s i m i l a r habitats as the species and reaching i t s maximum development on exposed slopes or mountain meadows where water slowly seeps through the extensive Philonotis mats. DISTRIBUTION: Endemic to North America and r e s t r i c t e d to the Coastal and Rocky Mountains from C a l i f o r n i a to the Aleutian Islands of Alaska (Map f i g . 70). Philonotis fontana var. americana was o r i g i n a l l y described by Dismier (1910) i n his r e v i s i o n of the North American species of Philonotis. His diagnosis and description i n French on page 22 c l e a r l y established the name as a subspecies of P. seriata Mitt. To comply with the 1906 Vienna Code of botanical nomenclature, Dismier provided a Latin description on page 35, however, th i s description was for P. americana sp. nov. Although the o r i g i n a l publication i s ambiguous, a l l of Dismier's annotation labels in NY read "P . americana Dism." which indicates that he 84 considered t h i s taxon a species rather than a subspecies. After the French description, Dismier c i t e d 41 specimens (syntypes) of which 27 are at NY. Of these 27, only 11 are beyond the range of v a r i a t i o n included i n the species concept of P. fontana. Considering the extreme v a r i a t i o n of the species, only robust " t y p i c a l " plants of var. americana can be i d e n t i f i e d with certainty. Persson (1954) has c l a r i f i e d the exclusion of P. seriata from North America, however, his opinion that ssp. americana i s c l o s e l y related to t h i s extra-North American species i s erroneous. Philonotis fontana var. americana can be recognized by i t s large size (5-10 cm) which i s the largest of the genus i n North America, the widely spaced erect spreading leaves with several p l i c a t i o n s on each side of the costa, and the very broadly ovate-lanceolate leaves that form a s p i r a l l i n g stem t i p . 85 Figure 61. Philonotis fontana var. amerioana 86 Philonotis fontana var. pumila (Turn.) B r i d . Bryol. Univ. 2:20. 1827. F i g . 62. Bartramia fontana var. pumilum Turn., Muse. Hib. 107. 1804. TYPE: " i n paludivus," (not seen). Philonotis fontana var. pumila f. longifolia Flow, in Grout, Moss F l . No. Amer. 2(3):176. 1935. TYPE: "Nevada, Baker 15 86," (Not in US as reported by Flowers). P. fontana var. pumila f. dimorphophylla Flow, in Grout, Moss F l . No. Amer. 2(3):176. 1935. nom. nud. P. tomentella Mol. in Lor., Moosstud. 170. 1864. syn. nov. P. fontana var. tomentella (Mol.) Jaeg., Ber. S. G a l l . Naturw. Ges. 1874-75:86. 1875. P. fontana ssp. tomentella (Mol.) Dism., B u l l . Soc. Bot. France 10. Mem. 17:26. 1910. P. fontana ssp. tomentella var. heterophylla Dism., B u l l . Soc. Bot. France 10. Mem. 17:26. 1910. TYPE: "B. C , Selkirk Mts. , Macoun 37 8," (not seen). P. fontana var. pumila f. heterophylla (Dism.) Flow, in Grout, Moss F l . No. Amer. 2(3):177. 1935. syn. nov. P. borealis (I. Hag.) Limpr., Laubm. Deutschl. 2:565. 1893. P. fontana var. borealis I. Hag., Kongl. Norske. Vidensk. Selsk. S k r i f t . 1888-90:13. 1889. TYPE: "In alpe Galdh a l t i t . ca. 1800 M. ad terram humidam 11/8, 1887," (not seen). 87 P. tomentella var. borealis (I. Hag.) Loeske., Verh. Bot. Ver. Brandenburg 47:338. 1906. (IV). P. fontana var. oompaota Schimp., Syn. Muse. ed. 2:520. 1876. P. tomentella var. compacta (Schimp.) Dism., Mem. Soc. Sc. Nat. Cherbourg 36:413. 1908. P. fontana var. alpina (Brid.) B r i d . , Bryol. Univ. 2:20. 1827. P. alpicola var. borealis (I. Hag.) Podp., Veroeff. Gepbot. Inst. Rubel Zurich 1:254. 1924. P. alpicola var. tomentella (Mol.) Torre et Sarnth., F l . T i r o l 5:412. 1904. Plants 1.0-6.0 cm high (mostly less than 5.0 cm), erect, yellowish to dark green, forming small mats containing 5.0-15.0 mm of green leafy stems over an accumulation of densely tomentose and closely arranged s t r a i g h t stems with dead brown leaves. Leaves 0.6-1.5 mm long by 0.3-0.7 mm wide, erect not crowded; + long lanceolate from a somewhat ovate concave base, st r a i g h t or f a l c a t e ; apex blunt i n very short leaves but usually acuminate with a percurrent or shortly excurrent costa occasionally becoming hyaline; margins revolute or rarely plane, doubly serrate with c e l l s p r o j ecting at both ends, less serrate at the base and singly serrate at the apex; lamina usually concave lacking p l i c a e , upper l e a f c e l l s oblong-rectangular (2-8:1), papillose at the lower c e l l ends on the abaxial surface, lower c e l l s s i m i l a r , 88 becoming proportionately larger but shorter (2-4:1). Dioicous, perigonia d i s c o i d (rosette shaped) bracts highly v a r i a b l e . Sporophyte small, seta 10.0-20.0 mm long; capsule spherical 1.0-2.0 mm; stomata 50-120 at base of capsule; peristome as for the genus; spores 18-24u, p a p i l l o s e . CHROMOSOME NUMBER: Not reported. HABITAT: Common i n a r c t i c or alpine regions on clay, s i l t , or rock, near creeks or seepage slopes growing with other bryophytes and vascular plants, or as pure sod. DISTRIBUTION: Circumpolar, extending south i n the alpine regions of the Rocky Mountains. Reported from a r c t i c Europe, Asia, Iceland and Greenland (Map f i g . 71). Philonotis fontana var. pumila i s a d i s t i n c t a r c t i c -alpine variety recognized by i t s small s i z e , s t r i c t , erect, and c l o s e l y packed stems clothed with abundant tomentum and small sporophytes. This taxon, treated i n the broad sense, has been variously recognized as P. fontana var. alpina, P. alpicola var., P. borealis, P. tomentella and a l l manner of combinations of these. Philonotis tomentella, sensu European workers, includes many of the small expressions of var. fontana. F e r t i l e plants are e a s i l y placed i n ei t h e r var. fontana or var. pumila depending on t h e i r size and ± erect habit, while s t e r i l e specimens beyond the d i s t r i b u t i o n a l l i m i t s of var. pumila should be included i n the species. 90 Philonotis yezoana Besch. et Card, in Card., B u l l . Soc. Bot. Geneve ser. 2, 1:123. 1909. TYPE: "Japon: Mori (n. 3505, ser. I ) ; Kominato (n. 42, ser. I ) ; Aomori (n. 406, ser. I; Kinashi, n. 3, 8, 46, 52); R i i s h i r i (n. 665); Hokodate (n. 1901, 2738); Osorezan (n. 2123); Ikaregaseki (n. 2669). Coree: Ouen-San (n. 591)," (not seen). F i g . 63. Plants 1.0-3.0 cm high, forming loose dark green t u f t s , strongly brown tomentose below, occasionally producing elongate deciduous propaguliferous shoots. Leaves 0.8-1.2 mm long by 0.4-0.7 mm wide, appearing lax, concave, and somewhat appressed, broadly ovate-lanceolate, s l i g h t l y acuminate; margins narrowly recurved near the base, serrate with single teeth or with low double teeth; costa stout, percurrent; upper c e l l s elongate (2-4:1) 6-9u wide, lamina c e l l s larger, cuboidal, rhomboidal, or i r r e g u l a r l y many sided (1-2:1) 8-14u wide, some leaves possessing borders 2-6 c e l l s wide of oblong or l i n e a r c e l l s , a l l basal c e l l s coarsely p a p i l l o s e over the middle of the lumen on both surfaces, less strongly on the abaxial surface, becoming progressively weaker toward the apex. Dioicous, perigonia d i s c o i d (forming a rosette). Seta 2.5-4.0 cm long; capsule spherical, 2.0-3.0 mm, stomata approximately 10 0, r e s t r i c t e d to the capsule base; peristome as for the genus; spores 19-24y, p a p i l l o s e . A l l of the North American material examined was s t e r i l e , therefore t h i s description was made from Japanese plants. CHROMOSOME NUMBER: Not reported. 91 HABITAT: Over rock i n shaded stream canyons, c l i f f s , or steep slopes that are wet by seepage water. DISTRIBUTION: Apparently rare, Vancouver Island, B r i t i s h Columbia mainland, C a l i f o r n i a , Montana, Vermont, Washington, and Japan (Map f i g . 72). These c i t a t i o n s represent the f i r s t report of P. yezoana from North America. Due to the dependence on u l e a f shape and other variable characters for i d e n t i f i c a t i o n these few c o l l e c t i o n s have been previously misinterpreted. I have examined Japanese specimens and they a l l compare well to the North American material, not only i n the characters mentioned i n the key but also i n the plants stature, dark green color, widely spaced stems forming loose t u r f s , and abundant tomentum. This rare species i s distinguished by having p a p i l l a e exactly i n the middle on almost a l l of the shortly rectangular or i r r e g u l a r l y cuboidal c e l l s , .and by having a border of two to s i x c e l l s that are two to three times as long as the lamina c e l l s . Ochi (1962) has combined t h i s taxon from Japan (along with several m i s i d e n t i f i e d specimens of P. fontana from North America) with P. fontana var. seriata (Mitt.) B r i d . I have examined the type specimen of P. seriata Mitt, i n NY, and when considering the s t a b i l i t y of p a p i l l a p o s i t i o n , marginal c e l l shape, and the presence of a border of elongate c e l l s , i t i s clear that Ochi's concept of P. fontana var. seriata i s too broad and that P. yezoana deserves s p e c i f i c status. 92 Figure 63. Philonotis yezoana 93 IX, PHYTOGEOGRAPHY OF PHILONOTIS The nine North American taxa of Philonotis display several noteworthy phytogeographic patterns. Those taxa with gemmiform perigonia (section Philonotula) are endemic to the Americas and reach t h e i r northern l i m i t of d i s t r i b u t i o n i n the southeastern United States. There i s a great d i v e r s i t y of species, including a considerable number of endemics (44), i n Central and South America, and i n the West Indies. Philonotis longiseta, glauoescens, and sphaerooarpa represent northern extensions from t h i s center of species d i v e r s i f i c a t i o n . Those taxa with d i s c o i d perigonia (section Philonotis) are H o l a r c t i c . Philonotis fontana i s the most widespread of a l l the North American species, and i s d i s t r i b u t e d almost throughout the H o l a r c t i c . Philonotis fontana var. pumila occurs i n the a r c t i c and extends south only i n alpine regions. Philonotis fontana var. americana i s the only endemic taxon i n North America, and i s d i s t r i b u t e d throughout the Rocky and Coastal Mountains. Philonotis oapillaris and P. marchica occur sporadically throughout Europe and adjacent regions while i n North America each shows a d i s t i n c t d i s t r i b u t i o n pattern; P. oapillaris i s r e s t r i c t e d to the P a c i f i c Coast and P. marchica i s widespread i n eastern North America and the southern Rocky Mountains. Philonotis yezoana i s apparently rare i n North America and i s disjunct from Japan. It i s d i f f i c u l t to say exactly where the North American taxa of the section Philonotis originated. However, the largest number of taxa occur i n A f r i c a and southern Asia (approximately 40 species) which may indicate another center of species d i v e r s i f i c a t i o n and the subsequent migration to North America. The a f f i n i t i e s of the North American species of Philonotis are with those of Eurasia (P. fontana, marchica, and capillaris), the West Indies, Central and South America (P. longiseta, glaucescens, and sphaerocarpa) , and Japan (P. yezoana) . 95 96 97 98 ' >fO i - j I M cO ao 30 Map f i g . 67 Philonotis marchioa (Hedw.) Bri d . 99 100 101 102 Map f i g . 71 Philonotis fontana var. pumila (Turn.) Brid. 103 104 X I . EXCLUDED TAXA Philonotis caloarea(B. S. G.) Schimp., C o r o l l . 86. 1856. Grout (19 35) , Lesquereux & James (1884), and Jennings (1951) have reported t h i s species from North America with reservations about i t s p o s i t i v e i d e n t i f i c a t i o n . Holzinger issued an e x s i c c a t i c o l l e c t i o n , reported by Bailey (1926), as No. 414 of his Musoi Acrocarpi. This c o l l e c t i o n consists of robust plants of P. fontana var. fontana. A l l other c o l l e c t i o n s i d e n t i f i e d as P. calcarea are referable to other North American taxa and do not compare at a l l to authentic European material. Philonotis leiophylla Kindb. in Mac, B u l l . Torrey Bot. Club 15:185. 1888. nom. nud. Philonotis mohriana (C. Mull.) Jaeg., Ber. S. G a l l . Naturw. Ges. 1873-74:90. 1875. P. mohvii C. Mull, ex Kindb., Enum. Bryin. Exot. 70. 1888. This plant, which belongs i n the genus Bveutelia, was co l l e c t e d i n Mexico and mistakenly credited to Louisiana as thoroughly explained by Steere (19 49). Philonotis pumila Kindb., Eur. No. Amer. Bryin. 2:328. 1897. TYPE: "Can. Ontario near B e l l e v i l l e , Macoun," (Lectotype S seen). The probable type was seen from Kindberg's herbarium. It i s apparently a species of Oncophorus, and c l e a r l y not a species of Philonotis. 105 Philonotis seriata Mitt., Jour. Linn. Soc. Bot. Suppl. 1:63. 1859. TYPE: "In Helvetia, Schleicher. In Britannia i n monte Ben-na-Bourd, W. Gardiner," (Holotype Gardiner NY seen). P. fontana ssp. seriata (Mitt.) Dix., Stud. Handb. B r i t . Moss. 294. 1896. P. fontana var. seriata (Mitt.) Kindb., Bih. K. Svensk. Vet. Ak. Handl. 7(9):255. 1883. There are no North American specimens of Philonotis that compare with the type of P. seriata. Many of the European and extra-European specimens also do not compare well with the type, which indicates that t h i s problematic taxon needs further study on a world wide basis. Philonotis wilsonii (B. S. G.) Mitt., Jour. Sinn. Soc. Bot. 7:153. 1863. This l i t t l e moss does occur i n eastern North America but belongs i n the genus Bartramidula. Philonotis falcata (Hook.) Mitt., Jour. Linn. Soc. Bot. Suppl. 1:62. 1859. Bartramia falcata Hook., Trans. Linn. Soc. London 9:317. 1808. I have seen P. falcata from Japan, Taiwan, the P h i l i p p i n e s , and India. I t i s very s i m i l a r to P. marchica as also suggested by Mitten (1859), Kabiersch (1937), and Ochi (1962). The only characters separating the two species are the somewhat larger size of both the gametophyte and sporophyte, and the arrangement of leaves i n v e r t i c a l rows i n P. falcata. It i s not always possible to d i s t i n g u i s h small and s t e r i l e specimens except on the basis of geographic d i s t r i b u t i o n . Philonotis falcata and P. marchica are probably vicarious i f not conspecific. 107 X I I . SYNONYMS Bartramia B. fontana = Philonotis fontana B. fontana var. pumilum - P. fontana var. pumila B. glauces cens = P. glauces cens B. longiseta = P. longiseta B. muehlenbergii - P. marchica B. radicalis = P. longiseta B. radicalis var. porteri = P. longiseta B. tenella = P. glaucescens B. uncinata = P. sphaerocarpa Mnium M. fontanum = P. fontana M. marchicum = P. marchica M. sphaericarpon = P. sphaerocarpa Philonotis P. acutiflora = P. fontana P. acuti folia = P. fontana P. adpressa = P. fontana P. alpicola var. borealis = P. fontana var. pumila P. alpicola var. tenuis = P. fontana P. alpicola var. tomentella = P. fontana var. pumila P. amerioana = P. fontana var. amerioana P. amerioana var. gracilescens = P. fontana P. amerioana var. torquata = P. fontana var. amerioana P. amerioana f. dimorphophylla = P. fontana var. amerioana P. amerioana f. Zaxa = P. fontana var. amerioana 10 8 P. arnellii = P. oapillaris P. borealis = P. fontana var. pumila P. brachy chypus = P. fontana P. caespitosa = P. fontana P. caespitosa var. adpressa = P. fontana P. caespitosa var. ampliretis = P. fontana P. caespitosa var. compacta Dism. = P. fontana P. caespitosa var. compacta Jaeg. = P. fontana P. caespitosa var. heterophylla = P. fontana P. caespitosa var. Zaxa = P. fontana P. calcarea f. Occidentalis = P. fontana P. fallax = P. fontana P. fontana var. adpressa =• P. fontana P. fontana ssp. adpressa = P. fontana P. fontana var. alpina = P. fontana var. pumila P. fontana var. ampliretis = P. fontana P. fontana var. aristinervis = P. fontana P. fontana f. a r i s t i n e r u i s = P. fontana P. fontana var. borealis = P. fontana var. pumila P. fontana var. brachyphylla = P. fontana P. fontana var. caespitosa = P. fontana P. fontana var. oapillaris = P. oapillaris P. fontana var. columbiae = P. fontana P. fontana var. compacta = P. fontana var. pumila P. fontana f. dimorphophylla = P. fontana P. fontana var. falcata = P. fontana P. fontana f. flowersii = P. fontana 109 P. fontana var. heterophylla = P. fontana p. fontana var. laxa = P. fontana p. fontana var. laxa f. tenuis = P. fontana p. fontana var. miaroblasta = P. fontana p. fontana var. microthamnia = P. fontana p. fontana var. pumila f. dimorphophylla = P. fontana var. pumila p. fontana var. pumila f. heterophylla = P. fontana var. pumila p. fontana var. pumila f. longifolia = P. fontana var. pumila p. fontana var. seriata f. oocidentalis = P. fontana p. fontana var. serrata = P. fontana p. fontana var. tomentella = P. fontana var. pumila p. fontana ssp. tomentella = P. fontana var. pumila p. fontana ssp. tomentella var. heterophylla = P. fontana var. pumila P. glabriuscula = P. marchica p. p. P. fontana P. glaucescens var. brevifolia = P. glaucescens P. glaucescens f. Zaxa = P. glauces cens P. glaucescens var. terrestris = P. tyZ-auceseens P. graoillima = P. glauces cens P. longiseta var. polygama = P. longiseta P. longiseta var. porteri = P. longiseta P. longiseta var. propagulaecaulis = P. longiseta P. longiseta f. propagulicaulis = P. longiseta P. macounii = P. capillaris 110 P. macounii var. torquata = P. fontana P. microcarpa = P. fontana P. muehlenbergii = P. marchica P. muehlenbergii var. tenella = P. glaucescens P. radicalis = P. longiseta P. seriata ssp. amerioana = P. fontana var. amerioana P. sphaerocarpa ssp. tenella var. terrestr-is = P. glaucescens P. sp ft a ere car-pa var. t e r r e s t r i s = P. sphaerocarpa P. subcapillaris = P. marchica P. tenella = P. glauces cens P. tenella var. coloradensis = P. fontana P. tomentella = P. fontana var. pumila P. tomentella var. borealis = P. fontana var. pumila P. tomentella f. brachycarpa = P. fontana P. tomentella var. compacta = P. fontana var. pumila P. torquata = P. fontana var. amerioana P. uncinata = P. sphaerocarpa P. uncinata var. glauces cens = P. glauces cens P. uncinata var. graoillima = P. gZaueeseens P. vancouveriensis = P. capillaris p. p. P. fontana I l l XIII, SUMMARY As with many other hygrophytic plants, Philonotis exhibits a great degree of environmentally induced phenotypic polymorphism. This phenotypic p l a s t i c i t y has resulted i n the description of many ecophenic modifications, which have been given formal taxonomic status. The present research was i n i t i a t e d to evaluate the taxonomic value of morphological characters of Philonotis. The experimental procedure adopted was to compare plants from the f i e l d with each other before and a f t e r c u l t u r i n g i n a uniform environment. Numerous c o l l e c t i o n s from various types of habitats produced plants that were morphologically d i f f e r e n t i n some characters. These plants were then cultured at the same time i n the same environment. Careful comparisons of these plants revealed that some characters are e a s i l y modified by the environment while other characters are stable within a taxon regardless of the environment. Also, some plants appearing quite d i f f e r e n t i n nature (from d i f f e r e n t habitats) appear i d e n t i c a l a f t e r c u l t u r i n g i n a uniform environment. This research shows, for the. North American species of Philonotis, that the p o s i t i o n of p a p i l l a e oh the l e a f c e l l s (top, middle, or bottom), nature of the marginal c e l l s (singly or doubly serrate), shape of the leaf (triangular or ovate-lanceolate), and the shape of the l e a f c e l l s (quadrate to elongate-rectangular), are stable characters not modified by the environment and therefore can be vised to support a 112 v a l i d c l a s s i f i c a t i o n . Leaf s i z e , arrangement, and amount of f a l c a t i o n ; the amount of curvature on the margin, l e a f apex shape, costa length, and p l i c a t i o n ; presence or absence of rhizo i d s , propagula, or peristome c i l i a ; and the shape and si z e of the perigonal bracts are a l l ecophenic characters too variable to have diagnostic value. Using the available experimental data, 29 names recognized by Flowers i n Grout (19 35) are shown to be based on ecophenic modification and may be reduced to synonymy. After c a r e f u l examination of type specimens and many extra-North American specimens, seven taxa may be excluded from the North American f l o r a . One taxon (P. yezoana) not previously recognized outside Japan i s newly reported from North America. Most species of Philonotis are dioicous with a chromosome number of n = 6. Philonotis longiseta i s monoicous, and the chromosome number of n = 12 i s a new report. After p l o t t i n g the d i s t r i b u t i o n of the. North American species i t can be shown that P. glauoescens, longiseta, and sphaerooarpa are predominantly i n the southeastern United States, and have t h e i r c l o s e s t a f f i n i t i e s with taxa of the West Indies, and Central and South America. In North America P. oapillaris i s r e s t r i c t e d to the P a c i f i c Coastal region while P. marchica i s much more widespread; both species occur i n Europe. Philonotis yezoana i s rare i n North America and also occurs i n Japan. Philonotis fontana i s very widespread i n North America and throughout the H o l a r c t i c . Philonotis fontana var. americana i s endemic to the P a c i f i c Coastal and Rocky Mountains, while P. fontana var. pumila i s a r c t i c and alpine. 114 XIV. EXSICCATI AND SELECTED SPECIMENS EXAMINED Philonotis glauoescens E x s i c c a t i Examined Grout, No. Amer. Musci Perf. 69 as P. uncinata (CAN, DPU, MICH, NO, SMU, TENN, TEX, UC, US), some specimens contain P. longiseta also, and some labels have inconsistent corrections. Renauld & Cardot, Musci Amer. Sept. 299 as P. tenella (CAN, MICH). Philonotis glauoes cens Selected Specimens Examined U. S. A. FLORIDA: Alachua Co., Devils Millhopper, Zales 2067, 2070, 2082 (UBC), Griffin 1750 (FLAS, GRO, LAF), Breen 1267 (FLAS, MICH, UT), McFarlin A36 58 as P. gracillima (CAN, FLAS, FSU), Anderson & Crum 13288, 13292 (CAN), G a i n e s v i l l e , Schornherst 2622 as P. gracillima (FSU). Citrus Co., Cryst a l River, Schallert 1933 as P. gracillima (CU) . Gadsden Co., Chattahoochee, Schornherst 1931 as P. gracillima (FSU). Hardee Co., Wauchula, McFarlin 22 Feb. 1938 (UT). Hernando Co., B r o o k s v i l l e , Zales 2041 (UBC). Leon Co., Tallahassee, Pursell 300 MF48 (DUKE), Breen 440 (UT), Schornherst 1607 as P. graoillima (FSU). Levy Co., Suwannee R. , Reese 734 (LAF), Manatee Springs St. Pk., Breen 3041 (FSU, NY), 1267 (UT), 3040 (FSU), 115 Fannin Springs, Schornherst 1713 as P. gracillima (FSU). Liberty Co., Rock B l u f f , McFarlin 1713 (UT), Rock Springs, Schallert M123 as P. sphaerooarpa (IA). Madison Co., Withlacoochee R., Schornherst 1354 as P. gracillima (FLAS). Manatee Co., Manatee R., Grout 22 Jan. 1941 (IA), March 1926 (UT), Ellenton, McFarlin 573 as P. tenella (FSU, MICH), Bradenton, Robinson 15 July 1953 as P. gracillima (US) . Marion Co., Juniper Springs, Conard 18 Feb. 1956 (IA), Lake Weir, Underwood 211 (IA, NY), Rainbow Springs, Breen 2874, 2879, 2898 (FSU), S i l v e r Springs, Breen 3363A (FSU). Orange Co., Apopka, Breen 3256 (FSU). Pasco Co., E l f e r s , Breen 2950 (FSU). Polk Co., Highlands City, Redfearn 431-50-55 (SMS)., Lake Wales, McFarlin 96 as P. tenella (UT), Winterhaven, McFarlin 205 (FSU). Seminole Co., Sanford, Rapp 125 (TENN), 22 Feb. 1921 (FSU). Suwannee Co., Ichatucknee Springs, Griffin 11574 as P. gracillima (FLAS), Redfearn 176-55 (SMS). GEORGIA: Monroe Co., High F a l l s St. Pk., Lampton 2420, 2423 (LAF), Stewart Co., Lumpkin, Grice 298-55 (SMS). LOUISIANA: Caldwell Parish, Columbia, Reese 9239 as P. gracillima (LAF). 116 Iberia Parish, New Iberia, Langlois 1893 as P. tenella (MICH). MISSISSIPPI: Wilkinson Co. , Woodville, Reese 11172 as P. graoillima (LAF). TEXAS: Anderson Co., Palestine, Whitehouse 22373 (SMU, UT). Burnet Co., Granite Mt., McVaugh 7651 (MICH, TENN, UBC, US). Comal Co., New Brumfels, Clover 24 Nov. 1933 as P. tenella (MICH), Whitehouse 18103a, 26213 as P. graoillima (SMU). Sabine Co., Milen, Reese 3790 (LAF). 117 Philonotis longiseta E x s i c c a t i Examined Grout, No. Amer. Musci Perf. 69 as P. uncinata, some specimens contain P. glauoescens (CAN, DPU, IA, MICH, US). Grout, No. Amer. Musci Perf. 264 as P. radicalis (DPU, IA, LAF, MICH, SMU, TENN, TEX, UBC, UC, US, UT). H a l l , Amer. Plains Flora 1870 as B. radicalis (F). Holzinger, Musci Aero. Bor.-Amer. 94 as P. radicalis (IA, L, MICH, OP, UC, US, WIS). Jermy, F l . Texensis 379 as B. radicalis (MO, US). Langlois, F l . Ludoviciana 442 as B. radicalis (US). Rapp, F l o r i d a Mosses 11 (OP). Renauld & Cardot, Musci Amer. Sept. 54 as B, radicalis (NY). Su l l i v a n t & Lesquereux, Musci Bor. Amer. 169 (ed. I 1856) as B. radicalis (MICH, MO). Su l l i v a n t & Lesquereux, Musci Bor. Amer. 254 (ed. II 1865) as B. radicalis (CAN, MICH). Philonotis longiseta Selected Specimens Examined U. S. A. ALABAMA: Baldwin Co., Spanish Fort, Zales & Jamieson 1904, 1909 (UBC) Conecuh Co., Evergreen, Webster & Wilbur 769 as P. glauoescens (MICH, TENN, UBC). Dale Co., Clayhatchee, Reese 2791 (LAF). Geneva Co., Hartford, Breil 1564 as P. sphaerooarpa (FLAS). Lee Co., Auburn, Earl & Baker 17 A p r i l 1897 as P. radicalis (NY), Letterman 133 as B. radicalis (MO). 1X8 Marion Co., Winfield, Harvill 7138 as var. porteri (UT). Mobile Co., Mobile, Mohr 5403 as P. radicalis (UT), Bel l i n g r a t h Gardens, Zales & Jamieson 1896, 1897 (UBC). Tuscaloosa Co., Warrior R., Harper 3298 as P. radicalis (DUKE, MO, NY, TENN, US). ARKANSAS: Faulkner Co., Cadron R., Hedfearn 25918 (SMS). Garland Co., Crystal Springs, Demaree 21992 (TENN), Hot Springs Nat. Pk., Demaree 22734 (US), Sharp 22734 (TENN). Merian Co., Buffalo R., Redfearn 15246 (SMS). Montgomery Co., Cedar Glade, Anderson 11761 (CAN, UBC). Newton Co., Ponca, Redfearn 12569 (MO, SMS, US). Saline Co., Owensville, Redfearn 24311 as P. graoillima (MICH, SMS). Scott Co., Waldron, Whitehouse 23357 (DUKE, SMU, US, UT). Stone Co., A l l i s o n , Anderson 11673, 11675, 11693, 11698 (UT). FLORIDA: Alachua Co., G a i n e s v i l l e , Murill 3 March 1938 (DUKE, UT), Zales, Jamieson S Griffin 2061-2065 (UBC). Calhoun Co., M i l l Creek, Wilson 3973 as P. graoillima (FSU, NY). Gadsden Co., Ochlocklonee R., Latina 52 (UBC). Hernando Co., Brooksville, Zales 2047, 2053, 2055 (UBC). Highlands Co., Highlands Hammock St. Pk., Griffin 11573 as P. glaucescens (FLAS). Homes Co., Ponce de Leon, Zales & Jamieson 1918, 1920 (UBC). Leon Co., Tallahassee, Sohornherst 1550 (FLAS, FSU, MICH), u 119 1016 (FSU, MICH), Zales & Jamieson 1931, 1939, 1940, 1941, 1948 (UBC). Levy Co., Suwannee R., Schornherst 1291 as P. glauoescens (MICH, US). Liberty Co., Torreya St. Pk., Zales 1930 (UBC), Redfearn 192-55 as P. glauoescens (SMS). Manatee Co., Manatee, McFarlin 433 as P. glauoescens (FSU). Marion Co., Eureka Springs, Schornherst 2026 as P. sphaerooarpa (FSU, MICH). Polk Co., Bartons, McFarlin 25 (FSU, MICH, UT). Seminole Co., Sanford, Rapp 125 (NY), Oviedo, Zales & Jamieson 2020, 2024, 2027, 2028, 2030 (UBC). Sumter Co., Wildwood, Robinson 13 July 1953 as P. glauoescens (US). Suwannee Co., Suwannee R., Schornherst 1390 (FLAS, FSU, MICH). Wakulla Co., Newport, Schornherst 1852 as P. glauoescens (NY) , 1853 (FSU) . GEORGIA: C a r r o l l Co., Yellow D i r t Creek, Zales <S Lampton 2129 (UBC), Whooping Creek, Zales & Lampton 2133 (UBC). Charlton Co., Folkston, Crosby 1868 (DUKE, MO). Fulton Co., Palmetto, Zales 2109, 2110, 2111, 2114 (UBC). Lowndes Co., Quarterman 3 (IA). Monroe Co., High F a l l s St. Pk., Zales 2086 (UBC). Rabun Co., T a l l u l a h F a l l s , Small 222 (DUKE, UBC). IOWA: Allamakee Co., Union City, Savage 1 June 1899 (IA). '-. 120 Marion Co., Red Rock, Coriard 14 May 1949 (IA). Muscatine Co., Wildcat Den, Savage 13 May 1899 (BRNM, IA, MICH, UT), Sweetland Creek, Shimek 8 May 1903 (DUKE, IA, MICH, TENN, US). KANSAS: Montgomery Co., El k ' C i t y , Smith 303 (NY). KENTUCKY: McCreary Co., Cumberland F a l l s St. Pk., Norris 129A (TENN). LOUISIANA: Avoyelles Parish, Mansura, Pennebaker 22 March 1940 (DUKE, UT) . Calcasieu Parish, Lake Charles, Shimek 29 Feb. 1936 (BRNM, DPU, DUKE, IA, MICH). Evangeline Parish, Easton, Reese 2662, 2665 (LAF). Iberia Parish, Weeks Island, Zales, Jamieson & Reese 1876, 1879, 1890, 1893, 1894 (UBC) . Lafayette Parish, Lafayette, Leon 30 March 19 31 (UT), Jan. 1931 (UBC), March 1933 (L, UBC), St. M a r t i n s v i l l e , Langlois March 1891 (US), Vermilion R., Zales 1861, 1863, 1866 (UBC). Ouachita Parish, Luna, Reese 8377 as P. glauoescens (LAF). Plaquemines Parish, Point a La Hache, Langlois 442 as B. radicalis (US). Rapides Parish, Zimmerman, MoWilliams <5 Davis 419 (LAF) . Sabine Parish, Many, Reese 67 71 (LAF). St. Landry Parish, Grand Coteau, Reese 7 867 (LAF). St. Martin Parish, Duchamp, Langlois 15 March 1892 as 1.21 B. radicalis (US). St. Mary Parish, Cote Blanche Is., Reese 7200 (LAF). West F e l i c i a n a Parish, Grant's Bayou, Reese 10802 as P. glauces cens (LAF). Winn Parish, Atlanta, Reese 6904 (LAF). MISSISSIPPI: Adams Co., Natchez, Shimek June 1898 (IA). Harrison Co., Wolf R. & Miss. Hwy. 53, Reese 3101 (LAF). Oktibbeha Co., Noxubee W i l d l i f e Refuge, Zales 1836 (UBC). Perry Co., Ragland H i l l s , Rogers 3676 as P. uncinata (FLAS). Wilkinson Co., Woodville, Reese 2557 (LAF). MISSOURI: Butler Co., Poplar B l u f f , Russell 78 (MO). Cedar Co., Stockton, Redfearn 12831 (SMS). Dade Co., Sylvania, Redfearn 8206 (SMS). Jefferson Co., P a c i f i c , Redfearn 20728 (SMS). St. Louis Co., Allenton, Nelson 2158, 1627 (DUKE, S-PA), 7646 (NY). Vernon Co., Nevada, Hall 1870 as B. radicalis (F). NORTH CAROLINA: Alleghany Co., Roaring Gap, Schallert 9872 as P. rigida (DUKE). Haywood Co., Lake Junaluska, Anderson 294 (DUKE). Jackson Co., E of Cherokee, Zales & Jamieson 2374 (UBC), Whitewater F a l l s , Sharp 24 Oct. 1959 (TENN). Jones Co., Trenton, Anderson 10792, 10794 as P. graoillima (DUKE). 122 Lee Co., Sanford, Anderson 20,554 (DUKE). McDowell Co., Woodlawn, Anderson & Jones 9671 (DUKE). Sampson Co., Newton Grove, Conard 17 March 1953 as P. sphaerooarpa (IA). Swain Co., Cherokee, Whitehouse 26821 (UT), Bryson City Anderson 819 (DUKE). OHIO: Hocking Co., Laurel Twp., Bartley & Pontius 412 (NY). Jackson Co., McCune, Bartley 609 (US). OKLAHOMA: Greer Co., Granite, Koch 5192 (MICH). LeFlore Co., Big Cedar, Redfearn 19786 (SMS), Ponteau, Redfearn 19592 (SMS), Short Mt., Redfearn 24723 (SMS). McCurtain Co., McCurtain Co. Game Preserve, R.edfearn 18720 (SMS). PENNSYLVANIA: LeHigh Co., Rockdale, Wolle Dec. 1874 (F) . Wyoming Co., Tunkhunnock, Glowenke 856 (IA). SOUTH CAROLINA: Charleston Co., Charleston, Smith A p r i l 1877 (US). Marion Co., Rains, Wonderly 867 (IA). Oconee Co., Sumter Nat. Forest, Wagner 9 June 1949 (TENN). Richland Co., Columbia, McCorkle B-98 (IA), 17 A p r i l 1950 (UT) . TENNESSEE: Blount Co., Abrams F a l l s , Sharp 341040 as f. polygama (TENN, UT) , Kinzel Springs, Sharp 34598 (F, MICH, TENN), 123 Rich Mt., 24 June 1934 (FSU). DeKalb Co., -Caney Fork Creek, Sharp 51T14 (UT). Monroe Co., T e l l i c o R., Sharp 1 May 1932 (DPU). Polk Co., Hiwassee R., Sharp 4911 (DPU, DUKE, MICH, TENN, UBC) . Sevier Co., Tremont, Grout & Sharp 2643 264a (LAF, UBC, US) . TEXAS: Bastrop Co., Bastrop, McAllister 19 A p r i l 19 36 (DUKE, SMU, TEX) . Brazos Co., White Creek, Parks 15 March 1947 (F, IA). Cherokee Co., Rusk, Whitehouse 22919a (SMU). Denton Co., Grapevine, Whitehouse 17918, 17943 (SMU, US). G i l l e s p i e Co., Grape Creek, Jermy 379 as B. radicalis (MO, US). Harrison Co., Caddo Lake St. Pk. , Whitehouse 27988 (SMU, US). Llano Co., Fredricksburg, McAllister 4 June 1919 (SMU, TEX), Austin, McAllister A p r i l 1938 (SMU, UT). Rusk Co., Mt. Enterprise, Whitehouse 20821a (SMU). San Jacinto Co., Pointblank, Reese 3597 (LAF, US). Tarrant Co., Hurst, Whitehouse 26292 (DUKE, SMU, US), 26531, 26538 (SMU). VIRGINIA: C a r r o l l Co., Fancy Gap, Gleason 47195 (MICH). Prince Edwards Co., Smith A p r i l 1878 (US). WEST VIRGINIA: Raleigh Co., Whitesville, Reed 74998 (US). 124 Philonotis sphaerocarpa E x s i c c a t i Examined Holzinger, Musci Aero. Bor.-Amer. 415 as P. uncinata (L, MICH, MO, UBC, UC, US, WIS). Husnot, PI. A n t i l l e s 147 as P. uncinata (L, WIS). Philonotis sphaerocarpa Selected Specimens Examined U. S. A. FLORIDA: Seminole Co., Sanford, Rapp. 128 as P. uncinata (OP). 125 Philonotis mavchiaa E x s i c c a t i Examined Austin, Musci Appal. 223 p. p. as B. fontana (CAN). Austin, Musci Appal. 225 as Bartramia, some specimens contain P. fontana (CAN, MICH, UBC, US, WIS). Holzinger, Musci Aero. Bor.-Amer. 471 as P. graoillima (CAN, L, MICH, MO, NY, OP, UBC, UC, US, WIS). Holzinger, Musci Aero. Bor.-Amer. et Eur. 543 as P. capillaris '(CAN, L, MICH, MO, NY, OP, UBC, UC, US, WIS). Macoun, Can. Mosses 173 (CAN) . Su l l i v a n t & Lesquereux, Musci Bor.-Amer. 168 (ed. I 1856) as B. muehlenbergii (CAN, MICH, MO, WIS). Su l l i v a n t & Lesquereux, Musci Bor.-Amer. 253 (ed. II 1865) as B. muehlenbergii (CAN, MICH). Philonotis marchica Selected Specimens Examined CANADA ALBERTA: Banff Nat. Pk., E Entrance, Crum & Schofield 5331 as P. fontana (CAN). Ghost Ranger Station, Bird & Glenn 11299a as P. fontana var. falcata (UAC). BRITISH COLUMBIA: Halcyon Hot Springs, MacFadden 4056, 19994 (CAN), 392 (UBC). NEW BRUNSWICK: Bass River, Fowler 1873 (L). NEWFOUNDLAND: Cow Head, St. Barbe, Tuomikoski 5021 (H). 126 NOVA SCOTIA: Anapolis Co., Kejimkujik Nat. Pk., Ireland 12511 (UBC). Inverness Co., Margaree Forks, Lamb 7012 as P. caespitosa (DPU, UBC) . ONTARIO: Bruce Co., Sauble Beach, Cain 1325 (UBC, UT). Carleton Co., Macoun 7 July 1900 as P. fontana (CAN). E l g i n Co., Sparta, James 35 as P. oapillaris (CAN). Essex Co., Leamington, Macoun 173 (CAN, MICH, US), 351 (CAN). Grey Co., Owen Sound, Crum 11614 as P. muehlenbergii (CAN). Hastings Co., B e l l e v i l l e , Macoun 5 June 1865 (CAN). Norfolk Co., Turkey Point, Cain 1330 (DUKE, IA, UT). Northumberland Co., Macoun 22 June 1864 as P. calcarea (CAN). Ontario Co., Lake Couchiching, Hand 818 (CAN, DUKE). Parry Sound Dist., Sundridge, White 249 as P. fontana (CAN). Wellington Co., Elora, Moxley 7 June 1933 (DUKE). QUEBEC: Gaspe, Fabius 4490 as P. oapillaris (DUKE, NY). U. S. A. ALABAMA: DeKalb Co., DeSoto St. Pk., Conard 5-8 June 1957 as P. sphaericarpa (IA). ARIZONA: Apache Co., Three Forks, Phillips & Mason 1572 (CAN, UT), as P. muehlenbergii (CAN). Cochise Co., Huachuca, Gooding 10329t 10345, 10030 (UBC, UT) . 127 Coconino Co., Oak Creek Canyon, Johnson 673 (UT), Marble Canyon, Clover & Jotter 2249 (CAN, MICH). Pima Co., Sycamore Canyon, Haskell 11 March 1945 as P. muehlenbergii (UT). Santa Cruz Co., White House Canyon, Bartram 101 as P. capillaris (CAN, DUKE, OP, US), Sycamore Canyon, Haring 3325 as P. muehlenbergii (UT). ARKANSAS: Boone Co., Harmony, Redfearn 25678 as P. muehlenbergii (SMS). Crawford Co., Natural Bridge, Redfearn 169 81 as P. muehlenbergii (SMS). Fulton Co., Mammoth Springs, Redfearn 15621 (SMS). Garland Co., Hot Springs, Scully 1101 (DUKE, IA, UT). Newton Co., Jasper, Anderson 11986 as P. muehlenbergii (DUKE), Ponca, Redfearn 12918 (SMS). Poinsette Co., Marked Tree, Demaree 17545 (US). Stone Co., A l l i s o n , Anderson 11616 as P.• muehlenbergii (DUKE, SMS). COLORADO: Gunnison Co., Gothic, Bunoe 92 as P. muehlenbergii (US). LaPlata Co., Hesperus, Pursell 3211 (CAN, DUKE, US). Summit Co., Tenmile Creek, Zales 3023 (UBC). FLORIDA: Jefferson Co., Monticello, Lightipe 117 (US). GEORGIA: Crisp Co., Cordele, Lamp ton 2278a as P. graoillima (LAF). Walker Co., Cloudland Canyon St. Pk., Anderson 12,973 as 128 P. uncinata (DUKE). IDAHO: Elmore Co., Atlanta, MaoFadden 19317 (CAN, DUKE, NY, UT). ILLINOIS: Lake Co., Lake B l u f f , Hill 22 Nov. 1904, 22 June 1905 (MICH). Menard Co., Wolf (US). INDIANA: Jasper Co., Remington, Welch 10116 (DPU). Jefferson Co., Hanover, Wolle 56 (DPU). Monroe Co., Bloomington, Welch 17420 (DPU). Putnam Co., Limedale, Baylor Spring 1904 as P. fontana (UC), Greencastle, Welch 10115 (DPU, MICH). Warren Co., L i t t l e Pine Creek, Ireland 5401 as P. gracillima (DPU, US) . IOWA: Allamakee Co., Quandahl, Welch 17270 as P. longiseta (DPU). Delaware Co., Delhi, Kurth Aug. 1946 (IA, SMU). Dubuque Co., Pine Hollow, Cavanagh 7290 (WJC). Emmet Co., Wellingford, Conard 7 Aug. 1946 (IA). VanBuren Co., Lucey-Keosauqua St. Pk., Conard 15 Oct. 1932 as P. longiseta (IA). Winneshisk Co., Bl u f f t o n , Conard 11 June 1949 (IA). KANSAS: Crawford Co., Farlington Lake, Gier 2376 (IA as P. longiseta, SMU) . Montgomery Co., Elk City, McGregar 2388 (IA). 129 KENTUCKY: Edmundsen Co., Brownsville, Price 28 as P. fontana (NY). Powell Co., Natural Bridge St. Pk., Conard 3 Sept. 1941 (IA) . LOUISIANA: West F e l i c i a n a Parish, Plettenberg, Reese 10486, 10491 as P. uncinata (LAF). MAINE: Cumberland Co., Peak's Is., Taylor 2875 (MICH). MARYLAND: Calvert Co., S c i e n t i s t ' s C l i f f , Chase 8412 as P. fontana (CAN). Garrett Co., Swallow F a l l s St. Pk., Worthley 27 Sept. 1964 (US) . MICHIGAN: Algar Co., Munsing, Conard 29 Aug. 1939 (IA). Barry Co., Lawrence Lake, Redfearn 22213 (MICH, SMS). Cheboygan Co., E of Burt Lake, Nichols 1920 (CAN, MICH). Chippewa Co., Scotty Bay Creek, no c o l l . c i t e d , 30 July .1920 (MICH) , Upper Tahquamenon F a l l s , Shetler 28 July 1958 (US). MINNESOTA: Blue Earth Co., Leiberg 393 as P. fontana (US). Hubbard Co., LaSalle Springs, Stein 58163 as P. muehlenbergii (UBC) . MISSISSIPPI: Clarke Co., Dunn F a l l s , Rogers 5762 as P. glaucescens (LAF). Clay Co., Hwy. 50 & Tombigbee R., Zales 1846 (UBC). 130 MISSOURI: Barry Co., Seligman, Redfearn 10269 (SMS). Benton Co., Avery, Redfearn 6922 as P. muehlenbergii (SMS). Boone Co., Ashland, Drew 3734 as P. oapillaris (MICH). Cedar Co., Bear Creek, Redfearn 12861 as P. muehlenbergii (SMS). Dade Co., Greenfield, Redfearn 6422 as P. muehlenbergii (SMS), Bona, Redfearn 7484, 6835 as P. muehlenbergii (SMS). Dallus Co., Celt, Redfearn 12788 (SMS). Douglas Co., Jackson's M i l l , Redfearn 8571 as P. muehlenbergii (SMS). Laclede Co., Sleeper, Redfearn 8262 (SMS). Madison Co., Minela Motte, Redfearn 159 58 as P. longiseta (CAN, DUKE, SMS). Newton Co., Boulder City, Redfearn 6676 (SMS). Phelps Co., James, Redfearn 7165 as P. muehlenbergii (SMS). Reynolds Co., Sec. 25, Gier 4609 as P. muehlenbergii (NY). Ripley Co., Buffalo Creek, Redfearn 6058 as P. muehlenbergii (SMS). Stoddard Co., Wapparello, Anderson 12567 as P. muehlenbergii (CAN, DUKE). Taney Co., Ozark Dam, Gier 462 as P. muehlenbergii (NY). Texas Co., A r r o l l , Redfearn 9 517 as P. muehlenbergii (SMS), Cabool, Redfearn 5182 as P. muehlenbergii (SMS). NEBRASKA: Cherry Co., Bog Valley, Walker 6 as P. fontana (DUKE). Thomas Co., Seneca, Kiener 20692 (CAN, MICH). 131 NEW MEXICO: San Miguel Co., Las Vegas, Arsene 18714 (US). NEW YORK: Fulton Co., Canada Lake, Ketchledge 555 (MICH, UBC, UT), Stoner Lake, Ketohledge 561 (MICH, UBC). St. Lawrence Co., Massena, Andrews 15 Sept. 1948 as P. capillaris (CU). Su l l i v a n Co., Willowemoc, Smith 37214 as P. fontana (MICH). Wayne Co., Wolcott, Cook July 1887 (NY). NORTH CAROLINA: Forsyth Co., Winston Salem, Chapman 2911 as P. graoillima (DUKE). Franklin Co., Riley, Beaven 677 as P. graoillima (DUKE), Weldon's M i l l , Beaven 563 as P. graoillima (DUKE). Haywood Co., Lake Junaluska, Watson 16 June 1928 as P. fontana (DUKE). Jackson Co., Whitewater R., Anderson 8669 as P. graoillima (DUKE). Macon Co., Coweeta, Webster 25 Nov. 1969 as P. muehlenbergii (TENN), Cullasaja Gorge, Schofield 9898 as P. graoillima (CAN, DUKE, UBC). Watauga Co., Blowing Rock, Zander 419 as P. graoillima (DUKE), Boon, Zales 2454 (UBC). OHIO: Athens Co., Athens, Robinson & Hall 25 May 1953 as P. longiseta (IA, US). Clark Co., S p r i n g f i e l d , Biddlecome 26 Sept. 1877 as 132 B. radicalis var. porteri (US). Lake Co., K i r t l a n d , Shipman 24 Oct. 1920 (NY). OKLAHOMA: Canadian Co., Devils Canyon, Little 13 as P. fontana (TENN). Cherokee Co., McSpadden F a l l s , Bird 3227 (UT). Murray Co., Turner F a l l s , Ikenberry 2080 as P. muehlenbergii (IA), Davis, Emig Aug. 1917 as P. calcarea (NY). Washington Co., Springdale, Anderson 12287 (CAN). PENNSYLVANIA: Allegheny Co., Douthitt, Jennings 26 A p r i l 1908 as P. fontana (MO, NY). Northampton Co. , Bethlehem, -Rau 17 May 19 47 as P. muehlenbergii (TENN), as P. fontana (NY). SOUTH CAROLINA: Lancaster Co., Taxahaw, Huntley 156 (DUKE). TENNESSEE: Davidson Co., Donelson, Clebsch 14595 as P. longiseta f. propagulicalis (TENN). Greene Co., Tusculum, Byers 10 Ap r i l 1949 as P. fontana (NY). Montgomery Co., C l a r k s v i l l e , Clebsch 808 (IA). Sevier Co., Laurel F a l l s , Zales 210 (UBC). TEXAS: Burnet Co., Burnet, Whitehouse 18502 as P. longiseta (SMU). Dallas Co., Dallas, Whitehouse 26302 (SMU, US). G i l l e s p i e Co., Palo Alto, Jermy 847 (MO). Mason Co., Mason, Harvill 2902 as P. longiseta (UT). Nacogdoches Co., Nacogdoches, Arnwere 24- (IA). 133 San Seba Co., Gorman F a l l s , Harvill 2915 as P. longiseta (UT) . UTAH: Kane Co., Escalante Ft. , Flowers 5624 (UT) , Kanab Canyon, Flowers 3248 (UT). S a l t Lake Co., Brighton, Flowers 572 (UT). San Juan Co., Temple of Music, Flowers 4822 (UT). VERMONT: Winsor Co., Norwich, Hutchinson 18 Oct. 1949 (DUKE, IA, SMU). VIRGINIA: Albemarle Co., Jones Run, Hermann 15173 as P. graoillima (CAN, DUKE, US). Botetourt Co., H o l l i n s , Patterson 3 July 1943 as P. muehlenbergii (UT). Greene Co., South R. F a l l s , Hermann 15132 as P. graoillima (US) , Ireland 3855 as P. glauces cens (US) . I s l e of Wight Co., Burwell's Bay, Correll 11487 (DUKE). James City Co., Williamsburg, Mukula 4647 (NY). Madison Co., Shenandoah Nat. Pk., Flowers 6426 (UT). Smyth Co., East Marion, Small 18 May 1892 as P. fontana (NY). WEST VIRGINIA: Mineral Co., Elk Garden, Ammons 1460 as P. muehlenbergii (UT). Mingo Co., Myrtle, Reed 70533 as P. muehlenbergii (US). Summers Co., Hinton, Morris 966 (NY, US, UT). WISCONSIN: Ashland Co., Copper F a l l s St. Pk., Hulbary 853 as P. longiseta (IA). Grant Co., Potosi, Cheney 11336 as P. fontana (WIS) WYOMING: Park Co., Clark Fork, Whitehouse 27497 (SMU). 135 Philonotis capillaris E x s i c c a t i Examined Holzinger, Musci Aero. Bor.-Amer. 440 (CAN, H, L, MICH, MO, NY, UBC, UC, US, WIS). Macoun, Can. Musci 152 as P. macounii (CAN, MICH, MO, NY, UAC, UBC, UC, US, WIS). Macoun, Can. Musci 416 as P. longiseta (MANCH). Renauld & Cardot, Musci Amer. Sept. 29 8 as P. macounii (CAN, MICH, NY). Philonotis opaillaris Selected Specimens Examined CANADA BRITISH COLUMBIA: B r i t t a n i a Beach, Schofield 12839 (UBC). Chilliwack, base of Mt. Cheam Zales & Schofield 1235 (UBC). Harrison Lake, Zales 2696 (UBC). Hope, Schofield 1669 8 (DUKE, UBC). Lund, Schofield & Boas 18157 (CAN, DUKE, UBC). Prince Rupert, Schofield & Boas 21490 (CAN, UBC). Skeena River, Schofield & Boas 20976 (UBC). Slocan City, MacFadden 9 June 192 7 (MO, UBC, UC). Taft, Schofield 29564 (UBC). Vancouver Island, Nootka Sound, Schofield & Halbert 40207 (UBC) . Queen Charlotte Islands, Graham Is., Skidegate Channel, Schofield 14001 (UBC), Moresby Is., Schofield 25266 (UBC). U. S. A. ALASKA: Juneau, Canby 487 (MO, NY, US). 136 Gold Creek Canyon, Cooley 31 July 1891 (NY). Kodiak, Trelease 1841 (MO, NY). Unalaska, Setchell 10 (UC). CALIFORNIA: Humboldt Co., Eel Rock, Koch 80 (MICH), Mad R., Schofield 28652 (UBC). Mendocino Co., Comptche, Zales 2899 (UBC), Fort Bragg, Zales 2893 (UBC), Leggett, Zales 2880, 2887 (UBC), Rockport, Schofield & Thomas 28 883 (CAN, DUKE, UBC). San Mateo Co., Lake San Andreas, Howe 136 (NY, UC, US). Santa Cruz Co., Big Basin Redwoods, Koch 2055 (MICH). Sonoma Co., Cazadero, Howe 14 March 1896 as P. fontana (NY) . T r i n i t y Co., Salyer, Tracy 16839 (UBC, UC). IDAHO: Kootenai Co., L i t t l e North Fork, Leiberg 172 (US), Tourlle R., Leiberg 96 (NY). OREGON: Lane Co., Leaburg, Leiberg 1697 (DUKE, US), Mapleton, Leiberg 1892 (US). Tillamook Co., Siuslaw Nat. Forest, Zales 2821 (UBC). WASHINGTON: King Co., North Bend, Ireland 5843 (CAN). Snohomish Co., Darrington, Schofield & Hermann 21828 (CAN, UBC), Gold Basin, Eyerdam 3007 (MICH, UBC), Mt. Pilchuck, Ireland 5590 (CAN), Stillaguamish R., Schofield, Ireland & Boas 18458 (CAN, UBC). 137 Philonotis fontana var. fontana E x s i c c a t i Examined Austin, Musci Appal. 223 as Bartramia p. p. P. marchica (CAN, MICH, NY, UBC, US, WIS), 224 as Bartramia (CAN, MICH, NY, UBC, US, WIS), 226 as Bartramia marchica (MICH, UBC, US, WIS). Backer, P a c i f i c Slope Bryo. 1586 (DUKE, MO, NY, UC, US, WIS), 766 (DUKE, MO, NY, UBC, UC, US, WIS). Bryhn, Mosses W. Minn. 22-26 June 1901 as P. alpicola (DUKE, MO, NY, US). Flowers, Mosses of Utah 8 (SMU). Frye, Moss Exs. 88 (CAN, MICH, NY, SMU, UBC). Grout, No. Amer. Musci Perf. 13 (CAN, MICH, NY, TEX, UC, US), 13c (CAN, DPU, LAF, MICH, NY, SMS, SMU, UBC). Grout, Hand-lens Mosses 46 (NY). Holzinger, Mosses N,-E. Minn. 15 Aug. 1902 as P. fontana var. caespitosa (BRNM, DUKE, NY, OP). Holzinger, Musci Aero. Bor-Amer. 70 (NY, OP, US), 316 as var. caespitosa (BRNM, CAN, L, MICH, MO, NY, OP, UC, US, WIS), 369 as var. seriata (CAN, L, MICH, MO, NY, UBC, UC, US, WIS), 414 as P. oalcarea (CU, L, MICH, MO, UBC, UC, US, WIS), 439 as P. amerioana (CAN, CU, L, MICH, MO, NY, UBC, UC, US, WIS), 441 as P. tomentella (CAN, MO, US), 441 as P. alpicola (CU, L, MICH, NY, UC, WIS), 517 as P. fallax (CAN, L, MO, NY, OP, UC, US, WIS). Holzinger, Musci Aero. Bor.-Amer. et Eur. 661 (CAN, L, MO, NY, UBC, UC, US, WIS). Leiberg, Musci Leibergiana 49 (CAN, DUKE, MICH, NY, UC, US, WIS) . 138 Macoun, Can. Mosses 170a as 'P. alpicola (CAN), 153 (UC, US). Nelson & Macbride, PI. Idaho 609 (MO, NY, US). Nelson & Holzinger, P i . Yellowstone Nat. Pk. 6062 (GRO, MO, NY, UC, US), 6110 as var. alpina (GRO, MO, NY, UC, US). Nelson & Holzinger, P i . Wyo. 1955 (MO, NY, UC, US), 6381 (MO, NY, US), 7718 (NY), 8770 (CU, MO, NY, UC, US). Sandberg, PI. Idaho 1115 (NY). Shaw, Selkirk F l . 983 as P. marchica (IA, L, MO). Small, Mosses So. U. S. 37 (CAN, DUKE, MO, NY, UBC, UC, US, WIS) . Su l l i v a n t , Musci Allegh. 123 as Bartramia (L, MICH, NY, US). Su l l i v a n t & Lesquereux, Musci Bor.-Amer. (ed. I 1856) 165 as Bartramia calcarea (MICH, MO, NY, WIS), 166 as Bartramia (MICH, MO, NY, WIS), 167 as Bartramia (MICH, MO, NY, WIS). Su l l i v a n t & Lesquereux, Musci Bor.-Amer. (ed. II 1865) 250 as Bartramia calcarea (CU, MICH, NY, WIS), 251 as Bartramia (CAN, MICH, NY, WIS), 252 as Bartramia (CAN, MICH, NY, WIS). Philonotis fontana var. fontana Selected Specimens Examined CANADA ALBERTA: Banff Nat. Pk., Kicking Horse Lake, Macoun 20 July 1885 (CAN), Lake Louise, Porsild & Breitung 17822 (CAN, DUKE). Cypress H i l l s Prov. Pk., Elkwater Bird 4454 (CAN, DUKE, NY, UAC) . Jasper Nat. Pk., Jasper, Crum 3657 (CAN, MICH). Waterton Lakes Nat. Pk., Goat Lake, Bird & Lakusta 16561 (CAN, UAC). BRITISH COLUMBIA: B a r k e r v i l l e , Schofield 39097 (CAN, DUKE). Brackendale, Macoun 193 (CAN). Cheam View, Fraser R., Schofield & Boas 17667 (CAN, TENN, UAC, US). Eagle Pass, Macoun 121 (CAN). Encampment, Svihla 42271 as var. pumila (CAN). Galiano Island, Salamanco Point, Schofield & Boas 17470 (CAN). Haines Hwy., Crum & Schofield 9194, 9213, 9615 (CAN). Hope, Williams 300 (CAN). Kootenay Lake, Lockheart Beach, Band 57-124 (CAN, DUKE). Li a r d Hot Springs, Porsild 22147, 22189 (CAN). L i l l o o e t , Noaxe Lake, Dill 1 Aug. 1961 (CAN, DUKE). Manning Prov. Pk., A l l i s o n Pass, Schofield 24764 (CAN). Macleod's Lake, Macoun 243 (CAN). New Denver, MacFadden 40 4 3 (CAN). New Westminster, Macoun 6 89 (CAN). Nicola R., Cougar Lake, Brinkman 10 July 1910 (CAN). Quesnel, Macoun 29 May 1875 (CAN). Rogers Pass, Selkirk Mts., Macoun 174 (CAN). Selkirk Mts., Armstrong, Wilson 70b (CAN). Slocan City, MacFadden 4054 (CAN). Vancouver Island, Elk Lake, Macoun 90 (CAN), Mt. Benson, Macoun 184 (CAN), Shawnigan Lake, Macoun 170a as P. alpicola (CAN), V i c t o r i a , Macoun 140 (CAN). Wells Gray Prov. Pk., Battle Mt., Ahti 13305, 14254 (CAN). 140 Whistler Mt., Schofield 37897 (CAN), Zales 1199, 2614 (UBC). Yoho Prov. Pk., Hector, Crum & Schofield 4499 as var. falcata (CAN, MICH, UBC). MANITOBA: Gillam, Kettle Rapids, Crum 7543 (CAN, UBC). Herb Lake, Wekusko Lake, Scoggan 104 (CAN, OP, SMU). Spruce Woods Prov. Pk., Melbourne, Bird 585 8 (CAN, MICH, NY, UAC). York Factory, Lower Hayes R., Ritchie 4229 (CAN). NEW BRUNSWICK: Albert Co., Fundy Nat. Pk., Ireland 11599 (DUKE, MICH, NFLD, UAC). Kent Co., Bass River, Fowler 18 June 1872 (L). V i c t o r i a Co., Grand F a l l s , Habeeb 939 (MICH). NEWFOUNDLAND: Avalon Peninsula, Colinet, Smith 427 (CAN). Battle Harbour, Waghorne 30 Aug. 1891 (CAN). Bonne Bay, Elkington Bill (NY). Cape Caribou, Grand Lake, Kallio 18 July 1963 (CAN). C h u r c h i l l F a l l s , Brassard 5032, 5238, 5487 as P. tomentella (NFLD) . Hampden, Tuomikoski 3089 (H). LaPoile, Tuomikoski 2066 (H). • Lejeune Lake & Abel Lake (between), Harper 3388 (CAN, US). Main Brook, Damman TD6027-4 (CAN). NOVA SCOTIA: Colchester Co., F a l l e i g h Lake, Prince 6098 (MICH, OP). 141 Inverness Co., Cape Breton Highlands Nat. Pk., Ireland 11868 (DUKE, MICH, NFLD, UAC, UBC). Kings Co., Black Hole, Schofield 10954 (NY). V i c t o r i a Co., Indian Brook, Schofield 6009 (NY, US). ONTARIO: Algoma D i s t . , Spanish M i l l s , White 242 (CAN), Montreal R., Lerstern 16 July 1960 (FLAS). Bruce Co., Hay Bay, Crum 11395 (CAN, MICH), Lions Head, Hand & Moxley 137 (CAN). Cockburn Island, Grassl 69 70 (MICH). Gray Co., Markdale, Crum 11254 (CAN, MICH). Kenora Dist., Nextor F a l l s , Hand 1159 (CAN). Muskoka Dist., Coopers F a l l s , Cain 1326 as P. caespitosa (IA, MICH, UAC). Niagra F a l l s , Drummonds Island, 15 May 1901 (CAN). Owen Sound, West H i l l , Moxley July 1926 (CAN). Parry Sound Dist., South R., Hand 843 (CAN). Peel Co., Orangeville, Cain 4956 (CAN, NO). Renfrew Co., Deep River, Crum 1 June 1955 (CAN). Simeoe Co., Sparrow Lake, Cain 1327 as P. amerioana (DUKE, IA, MICH, NO, NY, TENN, UAC, UBC). Thunder Day Dist., Black Fox Lake, Brodo 13806 (CAN, MICH), Ouimet Canyon, Garton 4477 (CAN), Nipigon, Macoun 26 June 1884 (CAN). PRINCE EDWARD ISLAND: Prince Co., Richmond, Ireland 10320 as P. caespitosa (MICH, UBC) . 142 QUEBEC: Gaspe Peninsula, Grosses-Roches, Jones 31953 (CAN). Gaspesian Prov. Pk., Mt. Albert, Crum & Williams 10582 (CAN, LAF, MICH, UBC), 10723 (CAN). Hudson Bay, Great Whale R., Brisson & Forest 20633, 21051, 21310 (CAN, NFLD, UBC). Levis, St. Henri, Masson & Gagnon 8597 (CAN). L u s k v i l l e , L u s k v i l l e F a l l s , Ireland & Ley 10025 (FLAS, MICH, UAC, UBC). Montmagny, Notre Dame du Rosaire, Masson & Gagnon 10090 (CAN). Richmond Gulf, Wiachewan Bay, Marr M371 (MICH). Riviere-du-Loup, Blouin 6550-1 (CAN). Terrebonne Co., Saint Jovite, LacMercier, Crum 9960 (CAN, LAF) . LaVerendrye Prov. Pk. (S o f ) , Crum & Williams 10184 (CAN). SASKATCHEWAN : Qu'Appelle Valley, Elbow Forest Preserve, DeVries 913 (CAN). YUKON TERRITORY: Dawson, Calder & Billard 3870 (NY). Dezadeash River, Alaska Hwy., Crum & Schofield 8658, 9508 (CAN). Mayo, Keno H i l l , Gillett & Calder 4378, 4266 (NY). U. S. A. ALABAMA: Randolph Co., Wedowee, Hermann 10034 (MO, US). Shelby Co., Montevallo, Blaokiston 6 A p r i l 1926 (NY). 143 ALASKA: Adak Isalnd, Shagak Bay, Jordal 2761 (CAN, MICH). Alaska Range, Mt. McKinley Nat. Pk., Smith 2202A (CAN, UC). Attu Island, Chicagof Harbor, Sohaack 691 (CAN, MICH, UBC, US). East Oumalik, Steere 15502 (CAN, MICH). Glacier Bay Nat. Monument, Muir I n l e t , Argus & Chunys 6545, 6547 (CAN, MICH). Juneau, Strong 55 as var. pumila (NY). Kenai Peninsula, Kenai R., Lutz 50-5, 50-6 (NY). Kodiak Island., Olga Bay, Looff E24S, E281 (CAN). Mt. McKinley Nat. Pk., Mt. Eielson, Weber & Viereck 10132 as P. tomentella (CAN, DUKE, SMS, SMU). Prince William Sound, Cooper 5 Sept. 1935 (NY). Skagway Quadrangle, Haines, Hermann 21799 (CAN). Unalaska Island, Maooun 303 (CAN). ARKANSAS: Conway Co., P e t i t Jean Mt., Anderson 11884 (DUKE). Crawford Co., Winfrey, Redfearn 21093 (SMS). Faulkner Co., M a r t i n v i l l e , Redfearn 19399 (SMS). Garland Co., Hot Springs, Russell 4 (NY). Newton Co., Ben Hur, Anderson 11933 as P. caespitosa (MICH). Polk Co., Mena, Anderson 11512 as P. muehlenbergii (DUKE, MICH). Stone Co., Ozark Nat. Forest, Redfearn 23164 (SMS). VanBuren Co., Shirley, Redfearn 16568 (SMS). Washington Co., Odell, Redfearn 16920 (MO, SMS). 144 White Co., Steprock, Redfearn 19427 (SMS). Y e l l Co., Dardanelle, Redfearn 23917 (SMS). ARIZONA: Cochise Co., Rucker Canyon, Pultz, Barrow, Gould & Phillips 2765 (NY). Coconino Co., Grand Canyon Nat. Pk., Raring & Bryant 3721 (NY), S of Jacob Lake, Zales 2982 (UBC). G i l a Co., Payson, Smith 9 May 1945 (NY). Graham Co., Mt. Graham, Pultz, Barrow & Phillips 3 Sept. 1944 (NY). Pima Co., Romero Canyon, Baring 3260, 3262 (NY). Santa Cruz Co., Baldy T r a i l , Bartram 172 as P. tomentella (CAN, DUKE, OP, US). CALIFORNIA: Butte Co., West Branch Campground, Kowalski 10 Oct. 1967 (CAN). Calaveras Co., Stanislaus R., Koch 1529 as var. pumila (NY). Eldorado Co., Upper Echo Lake, Beetle 4054 (NY). Fresno Co., Shaver Lake, Ikenberry 1602 (CAN). Humboldt Co., Grouse Mt., Tracy 14088% (UC). Inyo Co., Mt. Whitney, Baring 4523 (NY). Marin Co., Mt. Tamalpais, Koch 2291 (MICH). Mariposa Co., Yosemite F a l l s , Koch 1706 (MICH). Mono Co., Leevining Grade, Koch 17 79 (NY, UC). Napa Co., Napa-Monticello Rd., Koch 1366 (NY, UC, US). Placer Co., Donner Lake, MacFadden 16958 as var. pumila (CAN), Canyon Creek, MacFadden 16954 as var. pumila (CAN). 145 Plumas Co., Feather River Canyon , 27 A p r i l 1970 (CAN), LaPorte, Traverse 505 (CAN). Riverside Co., San Jacinto Mt., Hall 2388 (UC). San Bernardino Co., Snow Valley, Haririg 4515 (NY). Shasta Co., S t i l l w a t e r , Backer & Nutting 18 May 1894 (NY). Siskiyou Co., Mt. Shasta, Frye 10 July 1933 (SMU). Stanislaus Co., Oakdale, Zales 2922 (UBC). Tualumne Co., Sonora Pass, Howell 5 June 1954 (CAN, US). Tulare Co., E Fork Kaweah R., Koch 2168 (NY, US). COLORADO: Boulder Co., Boulder, Conard 41-214 (CAN). Chaffee Co., Monarch Pass, Zales 3014 (UBC). E l Paso Co., Manitou, Jewett 883 (NY). Grand Co., Rabbit Ears Pass, Schofield 7177 (CAN). Larimer Co., Fort C o l l i n s , Hermann 16968 (CAN, DUKE, US), Bear Lake, Zales 3058 (UBC). Mineral Co., Pagosa Peak, Baker 121 (NY)-. Ouray Co., Red Mt., Summit, Griffin 121 (FLAS). Park Co., Jefferson, Weber B-4391 (CAN, NY). P i t k i n Co., Aspen, Weber B-10912 (CAN, LAF), B-10898, 10899 (LAF). San Juan Co., E Lime Creek, Zales 3000s 3002 (UBC). Summit Co., Blue Lake, Weber & Anderson B-34256 as P. tomentella (CAN), Tenmile Creek, Zales 3027 (UBC). CONNECTICUT: New Haven Co., Carmel Mt., Eaton 5 May 1880 as Bartramia (NY) . 146 GEORGIA: C a r r o l l Co., Yellow D i r t Creek, Zales 2119-2122 (UBC). Dekalb Co., L i t t l e Stone Mt., Small 9577 3 9589 (NY, UBC). Rabun Co., T a l l u l a h F a l l s , Small 9523 (NY). Walker Co., Cloudland Canyon St. Pk., Zales 2147 (UBC). Whitefield Co., Rock Face Mt., Harper 279a (NY, US). IDAHO: Bonner Co., Lake Pend d ' O r e i l l e , Leiberg 49 (CAN, NY, US). Custer Co., Fourth of July Creek, MaoFadden 18737 (CAN, DUKE, MICH, NY, SMS). Elmore Co., Queens River, MaoFadden 18739 (CAN, MICH, NY, US). Idaho Co., Burgorf, Hermann 20291 as P. amerioana (NY). Kootenai Co., Leiberg 377. as. var. gracilescens (CAN, NY). Latah Co., Beal's Butte, Mueggler 8M-37 (CAN, DUKE). Shoshone Co., Marks Butte, Mahler 2313 (SMU). Teton Co., Vi c t o r , Payson 2181 (NY). ILLINOIS: Johnson Co., Go r e v i l l e , Redfearn 18739 (SMS). LaSalle Co., Starved Rock St. Pk., Voth & Richards 958 (DPU, NY). Pope Co., Burden F a l l s , Chase 11676 (CAN). INDIANA: Elkhart Co., Simonton Lake, Deam 54644 (DPU). Lagrange Co., Mongo, Welch 7635 (DPU, MICH). Tippecanoe Co., Lafayette, Welch 10132 as var. falcata (DPU). KENTUCKY-: Powell Co., Clay City, Wharton 124 (CAN, NY). 147 MAINE: Franklin Co., Rangeley, Oakes s. n. as P. caespitosa (IA, SMU) . Hancock Co., Mount Desert Island, Faxon & Band June 1890 as P. muehlenbergii (NY). Kennebec Co., Monmouth, Merrill 46 (NY). Oxford Co., Sumner, Parlin 12382 (NY). Penobscot Co., Lake Matagamon, Allard 6941 (NY, US). Piscataquis Co., Mt. Katahdin, Crane 562 (CAN). Sagadahoc Co., Georgetown, Friend 10505 (NY). Somerset Co., Skowhegan, Allen 6 July 1877 (NY). York Co., A l f r e d , Crane 10 (CAN). MARYLAND: D. C. Rock Creek Park, Maxon 6290 (NY). MASSACHUSETTS: Dukes Co., Nashawena Island, Northrop 5 July 1901 (NY). Essex Co., Amesbury, Huntington 10 March 1901 (NY). Suffolk Co., Blue H i l l , Faxon 197 (NY). MICHIGAN: Alger Co., Munising, Nichols 38 (MICH, NY). Barry Co., Yankee Springs, Mazzer 15 A p r i l 1962 (CAN, DUKE). Cheboygan Co., Iron Bridge, Ireland 4674 (CAN, US). Chippewa Co., Sugar Island, Crum July 1969 as P. capillaris (MICH). Delta Co., Big Bay de Noc, Gleason 2423 (DUKE, MICH, NY). Emmet Co., C e c i l Bay, Wynne 2674 (NY). Gratiot Co., Alma, Davis 22 Oct. 1892 (LAF). 148 Keweenaw Co., Copper Harbor, Hermann 16146 (CAN, US). Mackinac Co., Bois Blanc Island, Wynne 2581 (NY). Marquette Co., Marquette, Britton 83 (NY). Presque I s l e Co., Ocqueoc F a l l s , Crum 6 July 1947 (CAN). MINNESOTA: Big Stone Co., O r t o n v i l l e , Holzinger 22-26, 1901 as P. alpicola (NY). Cook Co., Grand Marais, Holzinger 316 as var. caespitosa (CAN). Lake Co., Two Harbors, Gleason 1940 as P. caespitosa (NY, UBC) . MISSOURI: Cedar Co., Cedar Springs, Ireland 3492 (CAN, US). Dade Co., Jordan Creek, Redfearn 6995 (SMS, US). Iron Co., Annapolis, Russell 63 (NY). Jasper Co., Neck, Palmer 3052 (CAN). Jefferson Co., P a c i f i c , Redfearn 25514 (SMS). Lincoln Co., Poley, Steyermark 104 (DUKE, NY). Madison Co., French M i l l s , Redfearn 20880 (SMS). Ripley Co., Pleasant Grove, Bush 10 Aug. 1899 (NY). Shannon Co., Eminence, Redfearn 8412 (FSU, LAF, SMS, TENN, US) . St. Charles Co., Schluersburg, Steyermark 156 (DUKE, NY). St. C l a i r Co., Blackjack, Redfearn 6853 (FSU, US). St. Genevieve Co., Pickle Springs, Redfearn 18386 as P. capillaris (MICH, SMS, TENN). 149 MONTANA: Flathead Co., McDonald F a l l s , Schofield 12192 (CAN, DUKE). G a l l a t i n Co., Bozeman, Chase 13858 (CAN). Glacier Co., Logan Pass, Hermann 18062 (CAN). Lake Co., Poison, Schofield 11740 (CAN, DUKE, TENN, US), Crane Creek Waterfall, ZaZ.es 3122 (UBC). Lincoln Co., Cabinet Mts., Schofield 11977 (CAN, DUKE). Madison Co., Pony, Rydberg <£ Bessey 1897 (NY). Missoula Co., Lolo Nat. Forest, Hermann 20183 (US). NEBRASKA: Garden Co., Oshkosh, Kiener 16227 as P. americana (MICH, TENN). NEVADA: Elko Co., Schoonover Creek, Lawton 2716 (CAN, DUKE, US). Ormsby Co., Kings Canyon, Backer 915 as P. aoutiflora (DUKE, NY). Washoe Co., Mt. Rose, Lawton 3081 (CAN, -DUKE, TENN, US). White Pine Co., Backer Creek, Lawton 2818 (CAN, DUKE,TENN, US) . NEW HAMPSHIRE: Coos Co., Gorhara, Ogden 2367 (CAN, DUKE, MICH, UAC). Crafton Co., Holderness, Faxton 192, 193 (NY). NEW JERSEY: Hunterdon Co., Lanibertville, Best 108 (CAN). Sussex Co., Montague Township, Nash 4 July 1909 (NY). NEW MEXICO: Cafron Co., G i l a Nat. Forest, Baad 1391 (MICH). 150 Dona Ana Co., Organ Mts.,•Wooton 1500 as P. amerioana (NY). Lincoln Co., Ruidosa Creek, Wooton 1516 as P. amerioana (CAN). Santa Fe Co., Santa Fe Canyon, Bartram 56 (CAN). NEW YORK: Essex Co., St. Huberts, Hermann 14804 (CAN, DUKE, NY). Green Co., Stony Clove, Smith 32687 (CAN, NY). Hamilton Co., Blue Mountain Lake, Hermann 15692 (CAN, US). Nassau Co., Massapequa, Cain 34 (NY). Rensselaer Co., Schaghticoke, Backer 2915 (NY). Rockland Co., Suffern, Austin June 1866 (NY). Steuben Co., Hornell, Smith & Ogden 13836 (NY). Tioga Co., Waverly, McNeilus 24 (FLAS). Ulster Co., Moose Lodge, Beats 661 (CAN). " Warren Co., Thurman, Smith 31049 (CAN, NY, US). NORTH CAROLINA: Alexander Co., Heddenite, Keever 490 (DUKE). Alleghany Co., Roaring Gap, Schallert 1195 (DUKE). Avery Co., Grandfather Mt., Anderson 12746 (CAN, DUKE, MICH). Davidson Co., Lexington, Anderson 9205 (DUKE). Durham Co., Durham, Blomquist 4535 (DUKE). Forsyth Co., Schallert 5-10-1930 (NY). Franklin Co., Bunn, Beaven 714 (DUKE). Jackson.Co., Oakland, Hermann 15340 (NY). Macon Co., Highlands, Ireland 2674, 2774, 2945 (CAN, US), B r i d a l V e i l F a l l s , Zales 2331, 2334, 2335 (UBC), Whiteside Mt., Zales 2350-2352 (UBC). McDowell Co., N Fork Catawba R., Zales 2452, 2453 (UBC). 151 Transylvania Co., Cashiers, Ireland 2580 (CAN, US). OHIO: F a i r f i e l d Co., Sugar Grove, Werner 3 Sept. 1892 (NY). Hocking Co., Cantwell C l i f f St. Pk., Redfearn 25278 (SMS). Jackson Co., Liberty Township, Bartley & Pontius 23 (NY). Lake Co., P a i n e s v i l l e , Werner 20 July 1892 (NY). OKLAHOMA: LeFlore Co., Cedar Creek Lake, Redfearn 24751 (SMS). Sequoyah Co., Black Fox Creek, Redfearn 24592 (SMS). OREGON: Clackamas Co., Timberline T r a i l , Thomas 317A (CAN). Benton Co., Mary's Peak, Lawrence 1381 (NY). Clamath Co., Buck Lake, Copeland 3648 as P. tomentella (NY). Mount Hood, Cloud Cap Inn, Miller 8428 (CAN). Multnomah Co., Multnomah F a l l s , Zales 2767 (UBC). Union Co., E l g i n , Sheldon 8731 (NY, US). PENNSYLVANIA: Bucks Co., Upper Black Eddy, Taylor 3614 (MICH). C l e a r f i e l d Co., Penfield, Moul 6029 as var. falcata (DUKE). Dauphin Co., Gratz, Altland 11 March 1938 (NY). McKean Co., Baliv a r River, Burnett 2556 (NY). Mercer Co., Grove City, Barbour 14 (BRNM, MICH, OP). Pike Co., Dingman's F a l l s , Wiegmann 10 Sept. 1944 (NY). Wayne Co., Five Mile Creek, Moul, s. n. (NY). RHODE ISLAND: Cumberland, Olney, s. n. (NY). SOUTH DAKOTA: Pine Ridge Reservation, Visher 2235 (NY). 152 TENNESSEE: Sevier Co., Charlie's Bunion, Schofield 8845 (CAN, DUKE), Clingmans Dome, Zales 2245 (UBC), Elkmont Campground, Zales 2161, 2167 (UBC), Laurel F a l l s , Zales 209, 2209 (UBC), Newfound Gap,.Zales 2313 (UBC). Unicoi Co., Unaka Springs, Sharp UM-691 (DUKE, MICH, TENN, US) . TEXAS: G i l l e s p i e Co., Coal Creek, Jermy 1890 as P. muehlenbergii (NY, SMU, US). J e f f Davis Co., Mt. Livermore, Hinckley as var. pumila 1136 (MICH, NY, TEX), 1137, 1147 (NY, TEX), 1148 (TEX). UTAH: Sal t Lake Co., Brighton, Darker 5763 (MICH). Washington Co., Zion Nat. Pk., Zales 2972 (UBC). VERMONT: Lamoille Co., Lake of Clouds, Grout 7-3-1896 (DUKE, NY). Rutland Co., Rutland, Tollison 10619 (NY). Windham Co., Bellows F a l l s , Githens 2735 (CAN, UAC). VIRGINIA: Albemarle Co., Jones Run F a l l s , Ireland 2198 (CAN, LAF, US), 2201, 2213 (CAN). Craig Co., Patterson R-673 (NY). Giles Co., Mountain Lake B i o l o g i c a l Station, Ireland 1128 (CAN, US). Lee Co., Cumberland Mt., Patterson SW-11 (CAN). Madison Co., Shenandoah Nat. Pk., Ireland 1903, 1918, 1928, 153 (CAN, US), 1927 (CAN, FSU, US). Page Co., Lewis F a l l s , Ireland 2147 (CAN, DUKE, US). Rockbridge Co., Cypress F a l l s , Carroll Nov. 1948 as P. caespitosa (NY). Shenandoah Co., Devils Hole Mt., Allard 4585, 4605, 6479 (CAN, DPU, MICH). Smyth Co., Bear Creek, Vail & Britton 7 June 1892 (NY). St. Paul Island, Macoun 1 Aug. 1892 (CAN). WASHINGTON: Chelan Co., Stevens Pass, Lawton 2399 (CAN), Zales 2627 (UBC) . King Co., W of Stevens Pass, Zales 2628 (UBC). K i t t i t a s Co., Lake Keechelus, Frye 88 (CAN). Lewis Co., Nisqually River, Schofield & Boas 22228 (CAN, DUKE). Olympic Nat. Pk. , Soldac Hot Springs, Schofield 19451 (CAN, DUKE). Pierce Co., Nisqually Glacier, Cowles 795 (CAN). Skamania Co., Skamania, Ireland & Simpson 9611 as var. falcata (CAN). Snohomish Co., Glacier Peak, Schofield & Hermann 21838 (CAN, DUKE). Whatcom Co., Mt. Baker, Schofield 16061 (CAN, DUKE), Zales 2567 (UBC). WEST VIRGINIA: Adams- Co., De l l s , Taylor 37 (NY). Pocahontas Co., Marlinton, Gray M-509 (OP). 154 Preston Co., Erwin, Gray 7512 (NO, NY). Tucker Co., Canaan Valley, Allard 10092 (CAN, SMU, US). WISCONSIN: Barron Co., Barron, Cheney 9981 (WIS). B a y f i e l d Co., Drummond, Cheney 4055 (WIS). Douglas Co., Amnicon F a l l s St. Pk., Schroeder M158 (LAF). Jackson Co., Black River F a l l s , 1859 (CAN, NY). Lincoln Co., Grandfather, Cheney 2612 (WIS). Milwaukee Co., Henderson 17 May 1884 (NY). V i l a s Co., Wisconsin River, Cheney 608 (WIS). Waukesha Co., Mukwonago, 10229 (WIS). Wood Co., Wisconsin Rapids, Chase 6470 (CAN). WYOMING: Albany Co., Brooklyn Lake, Porter 1427 (NY). Carbon Co., Ryan Park, Hermann 17179 as var. pumila (CAN, DUKE, NY, US). Park Co., Beartooth Butte, Conard 23 Aug. 1953 (CAN). Teton Co., Grand Teton Nat. Pk., Zales 3091 (UBC). 155 Philonotis fontana var. americana E x s i c c a t i Examined Baker, P a c i f i c Slope Bryo. 592 as P. alpicola (DUKE, MO, NY, UBC, UC, US, WIS). Al l e n , Mosses Cascade Mts. Wash. 52 as P. fontana (CAN, DUKE, DPU, MICH, NY, SMU, UBC, US). Holzinger, Musci Aero. Bor.-Amer. 70b (BRNM, CAN, L, MICH, NY, OP, UC, WIS). Philonotis fontana var. americana Selected Specimens Examined CANADA ALBERTA: Jasper Nat. Pk., Pyramid Mt., Ostafichuk 22 July 1970 as P. fontana (CAN). BRITISH COLUMBIA: Hastings, Macoun 4 May 1889 (CAN). Mt. Revelstoke Nat. Pk., Hamilton Cr. Brodo 7703 (UBC). Queen Charlotte Islands, Graham Island, Schofield 14 231 (DUKE). Vancouver Island, Port Hardy, Fort Rupert, Schofield & Williams 26778 (CAN, DUKE). SASKATCHEWAN: Cypress H i l l s Prov. Pk., Lone Pine Creek, Bird 4679 (NY, SMS, UAC). U. S. A. ALASKA: Adak Island, Shagak Bay, Jordal 2760 (DUKE, TENN, UBC, US). Kodiak Island, Rigg June 1913 as P. fontana var. seriata (NY). 156 Juneau, Meh.au May 1904 as P. seriata (NY) . Popof Island, Kinoaid 8-19 July 1899 (US). Unalaska, Coville & Kearney 1744 as P. fontana (NY). Yakobi Island, Shacklette 3937 (MICH). CALIFORNIA: Butte Co., J o n e s v i l l e , Copeland 1159 as P. tomentella (UBC). Del Norte Co., Bear Basin, Norris 9005 (DUKE, MICH). Lassen Co., Pine Creek, Baker & Nutting 12 July 1894 as P. fontana (NY). Madera Co., Yosemite Nat. Pk., Ikenberry 1088 (SMU, UAC). Modoc Co., Jess Valley, Grant 8258 (MICH). San Bernardino Co., San Bernardino Mts., Blesdale 66 as P. fontana (NY). Santa Cruz Co., Camp E l v i r s , Wagner & Ranzoni 2343 (MICH). Shasta Co., Lassen Nat. Pk., Koch 1933 (CAN, NO, NY, UC). Tehama Co., Lassen Nat. Pk., Koch 1945 (MICH, NY, UC). Tuolumne Co., Sonora Pass, Wiggins C-8 as P. fontana var. pumila (UBC) . IDAHO: Benewah Co., Potlatch, Alexander 7 July 1961 (SMU, UAC). Custer Co., Stanley, Hermann 19938 (US). Kootenai Co., mountain stream Leiberg 49 (NY). Latah Co., St. Joe Nat. Forest, Mahler 1860 (SMU, UBC). Valley Co., Lakefork Creek, Smith 5 July 1942 as P. fontana (CAN, DUKE, MICH, TENN, UBC, US). MONTANA: Glacier Nat. Pk., Ole Creek, Maquire & Piranin 5334 (UC). 157 Missoula Co., Lolo Hot Spring, Nixon 8 July 1951 (UBC). Yellowstone Nat. Pk., NW corner, Cain 4952 (NO, UAC, UBC). NEVADA: Washoe Co., Mt. Rose, Billings 1265 (DUKE, TENN). OREGON : Clackamas Co., Mt. Hood, Hermann 18738 (US). Douglas Co., Cry s t a l Springs, Redfearn 11659 (SMS). Hood River Co., Timberline T r a i l , Ireland 7114 as P. fontana var. -pumila (UBC). Lane Co., Blue R., Becking 24 July 1963 (UBC). Linn Co., E of Cascadia, Gilkey 6-43-A (IA). Multnomah Co., Larch Mt., Miller 8500 (OP). WASHINGTON: Chelan Co., Stevens Pass, Lawton W5479 as P. fontana (SMU). Grays Harbor Co., Quinalt R., Meyer 91 as P. fontana (DPU, SMU) . Pierce Co., Fairfax, Frye 20 May 1934 (CAN). Snohomish Co., Mt. Pilchuck, Eyerdam 3046 (UBC). Widbey Island, Gardner 1898 as P. fontana (US). WYOMING: Albany Co., Centennial, Koepper 5-10 July 1937 (DPU). Bighorn Mts., Paintrock Creek, Jack 4 Aug. 1900 as P. fontana (NY). Teton Co., Grand Teton Nat. Pk., Zales 3104, 3112 (UBC). Yellowstone Nat. Pk., Obsidian Creek, Castle 14 July 1921 (CU), NW corner on Hwy. 191, Zales 3121 (UBC). 158 Philonotis fontana var. -pumila Selected Specimens Examined CANADA ALBERTA: Banff Nat. Pk., Moraine Lake, Crum & Schofield 3946 as P. fontana (CAN, MICH, UAC, UBC), Lake Louise, Crum & Schofield 3833 as P. fontana (CAN, DUKE, MICH, UAC, UBC). Jasper Nat. Pk., Tonquin Valley, MaoFadden 20068 as P. fontana var. tomentella (CAN). BRITISH COLUMBIA: Sikanni R., N of Fort St. John, Correll 11930 (DUKE). LABRADOR: Port Burwell, Polunin 1082a-10 as P. tomentella (MICH). W Turnavik Island, Nutt 13 (US). MANITOBA: Fort C h u r c h i l l , Twin Lake H i l l , Crum & Schofield 6686 (CAN, NY, UBC). NORTHWEST TERRITORIES: Ba f f i n Island, A r c t i c Bay, Polunin 2590b as P. tomentella (MICH), Pangnirtung, Polunin 2643c as P. tomentella (MICH). Bathurst Island, Bracebridge Inl e t , Tener & Harington 37 5 (CAN). Belcher Island, Tukarak Island, Doutty 30 July 19 38 as P. caespitosa (NY). Bylot Island, Ec l i p s e Sound, Coombs 9 as P. tomentella (CAN). Devon Island, Basecamp Lowlands, Barrett 179, 199, 394, 399, (UBC), Truelove Lowland, Vitt 4388 (NFLD). Ellesmere Island, Tanquary Fiord, Brassard 1505, 1717, 1807a, 159 1884 (CAN, NY, UBC), 2531 (CAN, NFLD), Lake Hazen, Powell 8 (CAN, DUKE), E l l a Bay, Mohl 31 July 1965 (UBC). Great Bear Lake, Port Radium, Steeve 10032, 10124, 10230 as P. tomentella (CAN, MICH), 10230, 10503 (DUKE, MICH, • UBC). MacKenzie R. Delta, Eskimo Lakes, Scotter 9 583 as P. tomentella (UAC). Meighen Island, Kuo 23 Aug. 1968 as P. tomentella (CAN). Southampton Island, Coral Harbor, Ritchie 27 July 1954 as P. tomentella (CAN). QUEBEC: Hudson S t r a i t , Wakeham Bay, Polunin 1479a as P. fontana (MICH). U. S. A. ALASKA: Bering S t r a i t Dist., Ogotoruk Creek, Viereck & Bucknell 4618 as P. tomentella (CAN, UBC, US), Cape Dyer, Viereck & Bucknell 4185 as P. tomentella (CAN). Delta Junction, Donnelly Dome, Hermann 21142 (MICH). East Oumalik, Steere 15330, 15437 (CAN, MICH). Gubic, Chandler & C o l v i l l e R., Steere 16121 (NY). Hubbard Glacier, Disenchantment Bay, Coville & Kearney 1073 (NY) . Jago Lake, Jago River, Cantlon & Gills 1463C (CAN). Okpilak Valley, Okpilak Lake, Cantlon & Malcolm 0216 (CAN). Meade.River Camp, Steere 15671, 15685, 15703 (CAN, MICH). 160 COLORADO: Park Co., Mt. Lincoln, Weber 8829, 8824 as var. fontana (DUKE) . P i t k i n Co., Roaring Fork R., Weber B-10989 as P. fontana (CAN). MONTANA: Flathead Co., McDonald Creek, Hermann 18044 (CAN, NY, UBC). WYOMING: Park Co., Beartooth Lake, Welch 16794 (DPU). 161 Philonotis yezoana Selected Specimens Examined CANADA BRITISH COLUMBIA: Vancouver, Indian Arm, Zales & Schofield 1376 (UBC). Vancouver Island, 16 mi. W of Jordan River, Halbert & Price 2671, 2693 (UBC), Strathcona Prov. Pk., Zales 2505 (UBC). U. S. A. ALASKA: Sitka, Sharp, Sharp & Iwatsuki 5468a, 5492a (TENN). CALIFORNIA: Mariposa Co., Yosemite Nat. Pk., Vernal F a l l s , MaoFadden 17418 as P. fontana var. laxa f. tenuis (CAN), Tioga Rd., Flowers 5461 as P. caespitosa (UT). MONTANA: Flathead Co., Lake McDonald, Williams 446 as P. seriata (NY), Mt. L o t t i e Stanton, Holzinger & Blake 21 July 1898 (NY). Glacier Co., St. Marys Lake, Whitehouse 25497 as P. americana (DUKE, NY), Flowers 6127 as P. caespitosa (UT) . VERMONT: Mt. Mansfield, i n the Notch, 19 Aug. 1882, no c o l l . c i t e d (NY). The l a b e l c i t e s New Hampshire however Mt. Mansfield i s i n Vermont. WASHINGTON: Snohomish Co., Glacier Big Four, Schofield 20024 as P. fontana (UBC) . 162 XV. LITERATURE CITED Anderson, L. E. 1954. Hover's Solution as a Rapid Permanent Mounting Medium for Bryophytes. The Bryologist 57:242-244. & H. Crum. 19 58. Cytotaxonomic Studies on Mosses of the Canadian Rocky Mountains. B u l l . Nat. Mus. Can. Contr. to Bot. 160:1-89. Austin, C. F. 1877. New Musci. B u l l . Torrey Bot. Club 6(36):190. Bailey, J. W. 1926. Philonotis calcarea Schimp. i n Western Washington. The Bryologist 29:5-6. Bowering, M. L. 1966. The Autumn Meeting 1965. Trans. B r i t . Bryol. Soc. 5:211-212. B r i d e l , S. E. 1826-1827. Bryologia Universa. I I . L e i p z i g . B r i t t o n , E. G. 1911. Review of Dismier 1s Revision of Philonotis. The Bryologist 14:43-52. Brotherus, V. F. 1923. Die Laubmoose Fennoskandias. Flora Fennica I. Helsingfors. Bruch, P., W. P. Schimper & T. Giimbel. 1842. Genus Bartramia, Bryologia Europaea 4(12):315-326 p i . 1-11. Stuttgart. Buch, H. 1922. Die Scapanien Nordeuropas und S i b i r i e n s . I. Organographischer T e i l . Soc. S c i . Fenn. Comm. B i o l . 1(4):l-20. . 1928. Ibid. I I . Systematischer T e i l . Ibid. 3(1):1-177. Crum, H., W. C. Steere & L. E. Anderson. 1965. A L i s t of the Mosses of North America. The Bryologist 68:377-432. 163 Crundwell, A. C. 1970. I n f r a s p e c i f i c Categories i n Bryophyta. B i o l . Jour. Linn. Soc. 2(3):221-224. Davy de V i r v i l l e , A. 1927. L'action du M i l i e u sur les Mousses. Paris. Rev. Gen. de Bot. 40:156-173. Dismier, G. 1907a. Sur l a valeur specifique des epaississements i n t e l a m e l l a i r e s des dents peristomiales dans les especes du genre Philonotis. Rev. Bryol. 34:112-114. __. 1907b. Essai Monographique sur les Philonotis de France. Soc. Nat. S c i . Nat. & Math. Cherbourg Mem. 36:367-428. . 1910. Revision des Philonotis de l'Amerique. B u l l . Soc. Bot. France 10 Mem. 17:1-37. F i e l d , J. H. 1965. Possible Propaguliferous Branches i n Philonotis capillaris Lindb. Trans. B r i t . Bryol. Soc. 4:828. Florschutz, P. A. 1964. F l o r a of Suriname. Musci. VI(1) Leiden. Flowers, S. Mosses of Utah. Salt Lake City, Utah, (in prep.) Grout, A. J. 1935. Genus Philonotis by S. Flowers. Moss F l . No. Amer. 11:164-180. Newfane, Vermont. Hagerup, 0. 19 35. Zur P e r i o d i z i t a t im Laubwechsel der Moose. Det. Kgl. Danske Videnskabernes Selskab, B i o l . Meddelelser XI. 9:1-88. Hedwig, J. 1801. Species Muscorum Frondosorum. Leipzig. Heitz, E. 1942. Uber die Beziehung zwischen Polyploidie und Gemischtgeschlechtlichkeit bei Moosen. Arch. J u l i u s 164 K l a u s - S t i f t . Vererb., Sozialanth. u. Rassenhyg. 27:444-448. Hoagland, D. R. & D. I. Arnon. 1938. The Water Culture Methods for Growing Plants Without S o i l . C a l i f . Agr. Exp. Sta. C i r c . 374:1-32. Janzen, P. 1921. Die Blviten der Laubmoose: E i n Beitrag zur Kenntnis ihren ausseren und inneren Gestaltung. Hedwigia 62:163-281. Jennings, 0. E. 1951. A Manual of the Mosses of Western Pennsylvania and Adjacent Regions. Amer. Midland Nat., Monograph No. 6. Notre Dame, Indiana. Johansen, D. A. 19 40. Plant Microtechnique. New York. Kabiersch, W. 1938. Studien liber die ostasiatischen Arten ei n i g e r Laubmoosfamilien II (Bartramiaceae). Hedwigia 77:74-125. Kindberg, N. C. 189 7. European and North American Bryineae. Part 2. Linkoeping. . 1905. New North American Bryineae. Rev. Bryol. 32:33-39. . 1910. New Contributions to Canadian Bryology. The Ottawa N a t u r a l i s t 23:180-191. Lanjouw, J. & F. A. Stafleu. 1964. Index Herbariorum. Utrecht. Lesquereux, L. & T. P. James. 1884. Manual of the Mosses of North America. Boston. Lodge, E. 1960. Ef f e c t s of Certain C u l t i v a t i o n Treatments on the Morphology of Some B r i t i s h Species of Drepanocladus. Jour. Linn. Soc. Bot. 56:218-224. 165 Loeske, L. 1906. K r i t i s c h e Ubersicht der europaischen Philonoten. Hedwigia 45:195-212. Lowry, R. J. 1948. A Cytotaxonomic Study of the Genus Mnium. Mem. Torrey Bot. Club 20(2):l-42. Matteri, C. M. 1970. Morfologia del Protonema y Formacion de Yemas en Philonotis. Rev. Mus. Arg. Cs. Nat. (Bernardino Rivadaria) 3(8) :256-265 . Meusel, H. 19 35. Wuchsformen und Wuchstypen der europaischen Laubmoose. Nova Acta Leopoldina III 12:124-277. Mitten, W. 1859. Musci Indiae O r i e n t a l i s . Jour. Linn. Soc. Bot. Suppl. 1:1-171. Monkemeyer, W. 1927. Die Laubmoose Europas. i n L. Rabenhorst's Kryptogamen-Flora. IV. Leipzig. Nyholm, E. 1960. Moss Flora of Fennoscandia II (4):294-300. Lund. Ochi, H. 1962. Contributions to the Mosses of Bartramiaceae in Japan and the Adjacent Regions I. Nova Hedwigia IV(l-2):87-108. t l - 8 . . 1963. Ibid. I I . Ibid. V(1-2) :91-115. t7~24. Persson, H. 1954. Mosses of Alaska-Yukon. The Bryologist 57:189-217. Rau, E. A. & A. B. Hervey. 1880. Catalogue of North American Musci. Taunton. Schimper, W. P. 1855. Corollarium bryologia europaeae, conspectum diagnosticum familiarum. Stuttgartiae. Schwaegrichen, F. 1816. Species Muscorum Frondosorum. Supplementum Primum. Sectio Posterior. L e i p z i g . 166 Smith, A. J. E. & M. E. Newton. 1966. Chromosome Studies on Some B r i t i s h and I r i s h Mosses. I. Trans. B r i t . Bryol. Soc. 5:117-130. Steere, W. C. 1949. Some North American Mosses of Doubtful Antecedents. Rev. Bryol. et Lichen. 15 (1-2) : 24-27. Taylor, E. C. 1958. Residual Leaves from Brood-Branches of Philonotis caespitosa. The Bryologist 61:256-259. Vaarama, A. 1953. Some Chromosome Numbers of C a l i f o r n i a n and Finnish Moss Species. The Bryologist 56:169-177. . 1956. A Contribution to the Cytology of Some Mosses of the B r i t i s h I s l e s . The I r i s h Naturalist's Jour. 12 (2) :1-11. Yano, K. 1957. Cytological Studies on Japanese Mosses I. Mem. Fac. Ed. Niigata Univ. 6(3):1-31. Zastrow, E. 1934. Experimentelle Studien uber die Anpassung von Wasser und Sumpfmoosen. Jena. 

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