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Hepaticae and Anthocerotae of southwestern British Columbia Godfrey, Judith Louise Dean 1977

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THE HEPATICAE AND ANTHOGEROTAE OF SOUTHWESTERN BRITISH COLUMBIA by JUDITH LOUISE DEAN GODFREY B.A., Hiram College, I969 M.S., University of Minnesota, 1971 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of Botany We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1977 © Judith Louise-Dean Godfrey, 1977 In p r e s e n t i n g 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 requ i rement s f o r an advanced degree at the U n i v e r s i t y o f B r i t i s h Co lumb ia , I ag ree 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 r e f e r e n c e and s tudy . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f 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 g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s . It i s u n d e r s t o o d that c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i thout my w r i t t e n p e r m i s s i o n . Department o f B o t a n y  The U n i v e r s i t y o f B r i t i s h Co lumbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date September 15, 1977 1 It commonly chances that I make my most interesting botanical discoveries when I £am3 in a thrilled and expectant mood, perhaps wading in some remote swamp where I have just found something novel and feel more than usually remote from the town. Or some rare plant which for some reason has occupied a strangely prominent place in my thoughts for some time will present itself. My expectation ripens to discovery. I am prepared for strange things. — H. D. Thoreau Journal, September 2, I856 Schofieldia monticola Godfrey i i i ABSTRACT The Hepaticae of western North America, in contrast to eastern North America and many other parts of the world, are poorly known. This present flora for southwestern British Columbia, Canada, and part of northwestern Washington State, U.S.A., is intended to partially alleviate this problem. The geographical area covered in this study offers great diversity in topography, climate, and vegetation. The region includes the wet Pacific coast of western Vancouver Island, mesic forests and floodplains of low elevations, the dry area around southern Georgia Strait, subalpine forests and meadows of the Coast Range and North Cascade Mountains, alpine soil and rock substrates at highest elevations below summer snowline or glaciers, and dry habitats in the rainshadow northeast of the mountains. The material presented is based on direct study of approximately 4000 personal and 1800 herbarium collections of hepatics. A total of 166 species and two additional varieties, belonging to 64 genera in 37 families, are treated in keys and concise descriptions, accompanied by ecological and phytogeographical information. Additional discussion on taxonomic and systematic difficulties is given where pertinent. Five hepatics new to science were discovered during this study, with Schofieldia monticola Godfr. having been recently described. Hepatics collected which had not been reported previously from continental North America include Eremonotus myriocarpus (Carring.) Lindb. et Pears, in Pears., Jungermannia hattoriana (Amak.) Amak., Marsupella condensata (Angstr. inHartm.) Schiffn., and Nardia .japonica Steph. Hepatics belonging to, or similar to, Lophozia elongata Steph. and Marsupella  adusta (Nees emend. Limpr.) Spruce were collected. Hepatics new to British Columbia include Lophozia ventricosa var. silvicola (Buch) Jones, iv Riccia fros t i i Aust., Scapania gymnostomophila Kaal., and S. paludlcola Loeske et K. Mull, in K. Mull. For the fi r s t time, fasciculate gemmae were discovered in Ghandonanthus Mitt., and inflated lobules were found in Porella navicularis (Lehm. et Lindenb.) Lindb. Based on specimens exajnined from the study area, Bazzania ambigua (Lindenb.) Trev., Odontoschisma gibbsiae Evans, and Plagiochlla satoi Hatt. were treated as synonyms of other species in this flora. Systematic problems requiring detailed future study were encountered in particular in Bazzania S. Gray and Galypogeia Raddi emend. Nees. Regional distribution-maps depicting a l l known points of occurrence and general elevation are presented for a l l hepatics discussed in this flora. Four general categories of distribution were demonstrated by a comparison of the spatial patterns: l) distributions in moist, humid climates influenced to varying degrees by the Pacific Ocean (18$ of the total flora), 2) high elevations in the Coast and Cascade Mountains (2k%), 3) dry climates in rainshadow areas (16$), and 4) various types of widespread distributions (33$)• The remaining 9$ of the species are rare or infrequent and cannot be assigned to any particular category. The hepatic taxa were assigned to lb phytogeographical elements based on total worldwide distributions. Approximately 50$ of the hepatics have essentially circumboreal distributions. This figure includes taxa missing from eastern Asia. Ten percent of the hepatics are endemic to western North America. Of the hepatics demonstrating disjunct, or discontinuous distributions, 16$ have European affinities, 9$» eastern Asian affinities, and 11$, affinities with both regions. Approximately 11$ of the hepatics treated in this flora have bipolar disjunctions. In an attempt to reconstruct some events in the development of the modem hepatic flora of southwestern British Columbia, possible Pleistocene refugia and their effect on hepatic populations are discussed. This flora will provide a manual for the Hepaticae and Anthocerotae of southwestern British Columbia, and will serve as a preliminary guide to these plants in the North American Pacific Northwest. v i TABLE OF CONTENTS PAGE PART I: INTRODUCTION AND METHODS 1 INTRODUCTION 2 HISTORY OF HEPATICOLOGICAL STUDIES IN SOUTHWESTERN BRITISH COLUMBIA . 4 LOCATION OF THE STUDY AREA 8 CLIMATE OF THE STUDY AREA 11 PHYSIOGRAPHY AND GEOLOGY OF THE STUDY AREA 14 General Features and History 14 Pleistocene Geology 17 Volcanism and Alpine Glaciers 19 VEGETATION OF THE STUDY AREA 22 Coastal Douglas F i r Zone 25 Coastal Western Hemlock Zone 26 Subalpine Mountain Hemlock Zone 28 Forest Subzone 28 Parkland Subzone 29 Subalpine Meadows 29 Alpine Zone 30 Engelmann Spruce - Subalpine F i r Zone 32 Interior Douglas F i r Zone 33 Ponderosa Pine - Bunchgrass Zone 33 METHODS 35 General Methods 35 Description and Measurement 37 PART II: THE HEPATIC FLORA OF SOUTHWESTERN BRITISH COLUMBIA 40 FORMAT 41 System of Classification 43 General References 44 KEYS TO THE HEPATICAE AND ANTHOCEROTAE OF SOUTHWESTERN BRITISH COLUMBIA 45 CLASS HEP ATI CAE 60 Subclass Jungermanniae 60 Order Calobryales 60 Family Takakiaceae 60 Family Haplomitriaceae 62 Order Jungermanniales 63 Family Herbertaceae . 63 Family Blepharostomaceae 65 v i i PAGE Family Ptilidiaceae . . . 67 Family Lepidoziaceae 70 Family Calypogeiaceae 84 Family Jungermanniaceae . . . 96 Subfamily Lophozioideae . . 96 Subfamily Jamesonielloideae 128 Subfamily Jungermannioideae . 129 Subfamily Mylioideae 146 Family Gyrothyraceae 147 Family Gymnomitriaceae 148 Family Scapaniaceae 158 Family Antheliaceae 176 Family Cephaloziaceae 178 Subfamily Hygrobielloideae 179 Subfamily Gephalozioideae 179 Subfamily Odontoschismatoideae 188 Family Cephaloziellaceae 191 Family Geocalycaceae 197 Family Plagiochilaceae 202 Family Radulaceae 210 Family Pleuroziaceae 213 Family Porellaceae . . . . . . . . . 215 Family Frullaniaceae 219 Family Lejeuneaceae 225 Order Metzgeriales 226 Family. Fossombroniaceae 226 Family Blasiaceae 228 Family Dilaenaceae 228 Family Aneuraceae 233 Family Metzgeriaceae 237 Subclass Marchantiae . 239 Order Sphaerocarpales 239 Family Sphaerocarpaceae 239 Order Mar chant i ales 240 Family Targioniaceae . . . . 240 Family Grimaldiaceae 241 Family Gonocephalaceae 243 Family Lunulariaceae 244 Family Sauteriaceae 245 Family Marchantiaceae 246 Family Ricciaceae 247 CLASS ANTH0CER0TAE 251 Order Anthocerotales . . . . . 252 Family Anthocerotaceae ' 252 PART III: DISCUSSION 256 HEPATICS OF PARTICULAR INTEREST IN SOUTHWESTERN BRITISH COLUMBIA . . 257 New Records 257 Notes on Systematic Problems 260 v i i i PAGE NOTES ON HEPATICAE COLLECTED BY JOHN MACOUN IN SOUTHWESTERN BRITISH COLUMBIA 264 ECOLOGY AND REGIONAL DISTRIBUTION OF HEPATICS IN SOUTHWESTERN BRITISH COLUMBIA . . 272 PHYTOGEOGRAPHICAL RELATIONSHIPS OF THE HEPATICAE OF SOUTHWESTERN BRITISH COLUMBIA 292 Introduction 292 Floristic Elements 299 Continuous Distributions 299 Discontinuous, or Disjunct, Distributions . . . . 31k DEVELOPMENT OF THE MODERN HEPATIC FLORA IN SOUTHWESTERN BRITISH COLUMBIA 329 CONCLUSIONS . 3 3 ^ LITERATURE CITED 339 PART IV: APPENDICES 364 Appendix A. A checklist of the Hepaticae and Anthocerotae of southwestern British Columbia 365 Appendix B. World and regional distributions of the Hepaticae and Anthocerotae of southwestern British Columbia . . 370 Appendix C. Maps depicting regional distributions of the known Hepaticae and Anthocerotae in southwestern British Columbia, Canada, and portions of adjacent Washington State, U.S.A. . . . 376 Appendix D. Hepatics collected in southwestern British Columbia by John Macoun 419 Appendix E. Sample field-data sheet for use in collecting hepatics . 421 Appendix F. Approximate relative size categories for leafy and thallose hepatics . . . 422 Appendix G. A guide to features important in identification of Hepaticae 423 Appendix H. Glossary 427 Appendix I. Addendum: Nardia breidleri (Limpr.) Lindb 432 ix LIST OF TABLES TABLE PAGE I Simplified outline of geological history in southwestern British Columbia 18 II Late Pleistocene history of southwestern British Columbia and northwestern Washington 20 III Comparison of three species of Calypogeia in North America and Europe 86 IV A continuum of features displayed by plants in the Plagiochila asplenioides species-complex . . 206 V Chronological sequence of hepatic collections made by John Macoun and his party during April and May, 1889, in southwestern British Columbia, based on specimens in the collection of the National Herbarium, Ottawa 269 VT Some characteristic hepatics for the biogeoclimatic zones, or portions thereof, in southwestern British Columbia 289 VII Floristic elements represented among the Hepaticae of southwestern British Columbia 298 VIII Geologic time-scale since the Cretaceous Period 302 X LIST OF FIGURES FIGURE PAGE 1 Reference map for the study area 9 2 Annual precipitation in southwestern British Columbia 12 3 Accumulated annual temperature in southwestern British Columbia: Day-Degrees over 6° C 13 4 Physiographic subdivisions and generalized geology of southwestern British Columbia 15 5 Major zones of vegetation in southwestern British Columbia and northwestern Washington State 24 xi ACKNOWLEDGEMENTS I am very grateful to my graduate advisor Dr. Wilfred B. Schofield for his enthusiastic support of my hepaticological studies and for good advice during the field work and writing of this thesis. I thank Drs. T. Amakawa, S. Hattori, H. Inoue, and N. Kitagawa for providing much helpful advice and identifying or verifying various hepatic specimens during this study. I am grateful to Dr. R. M. Schuster for sending comments and valuable advice regarding Schofieldia. Field work was funded by the National Research Council of Canada through a grant to W. B. Schofield. I thank John Pinder-Moss for making possible the trip to Meager Creek, and Dr. Charles Hebant, for the 1974 trip to Jervis Inlet. Additional support for my graduate studies was gratefully received from the University of British Columbia as a Graduate Fellowship (1975-1977) and a Summer Research Scholarship (1973). I thank the National Herbarium, Ottawa, for making available for study the J. Macoun collection of hepatics. I am indebted to my parents, James W. and Clarice Dean for their encouragement and support throughout the 24 years of my academic education, and for sending me to Hiram College, where my interest in botany grew under the influence of Dr. Dwight H. Berg. Most of a l l , I thank my husband, Dr. Geoffrey A. Godfrey, for accompanying and assisting me in the field, participating in hepaticological discussions and inquiries, and for providing moral and other support throughout this study. Geoff's excellent abilities in the field in general, and now in finding Haplomitrium and other rare or new hepatics, are greatly appreciated. 1 PART Ii INTRODUCTION AND METHODS 2 INTRODUCTION Hepaticological research in western North America has been generally sporadic when compared with work done on hepatics in eastern North America and many other parts of the world. Because of this situation the Hepaticae of the area are poorly understood. This study was undertaken to increase the knowledge on hepatics in the Pacific Northwest, and to provide modern keys to species occurring within a portion of the region. New collections of hepatics were made from a variety of habitats, from sea-level to alpine elevations, in southwestern British Columbia, Canada, and portions of adjacent northern Washington State, U.S.A. Inclusion of climatically and topographically diverse sites in the study area makes this flora applicable over a more extensive region in the Pacific Northwest. From the examination of living plants collected, and also of specimens already contained in the Herbarium of the University of British Columbia, the fl o r i s t i c base of this study was established. In order to better understand the nature and relationships of the known hepatic flora of southwestern British Columbia and northwestern Washington, additional information was sought regarding morphological features of the gametophyte and sporophyte of individual taxa, environmental variation, phenology of sporophyte production, habitat and substrate conditions, and associated hepatics occurring with each species. These data are organized into keys and brief descriptions. In subsequent chapters the Hepaticae of southwestern British Columbia are discussed in terms of new species records; inherent taxonomic and systematic problems; ecological associations of the species; local distributions of individual hepatics; phytogeography of the hepatics on a worldwide basis; and development of the modern hepatic f l o r a i n the region. This flora, with i t s accompanying keys, w i l l serve as a basic manual to the hepatics of southwestern British Columbia, as well as a preliminary guide to liverworts of adjacent areas of the Pacific Northwest. It w i l l also provide an indication of which taxa require additional investigations in greater detail. 4 HISTORY OF HEPATICOLOGICAL STUDIES IN SOUTHWESTERN BRITISH COLUMBIA Studies of hepatics began in southwestern B r i t i s h Columbia in the late 1800's, but accumulation of knowledge has been sporadic since that time. Pearson's (1890) L i s t of Canadian Hepaticae included 55 species from B r i t i s h Columbia. Cooley (I892) reported plants collected i n Alaska and Nanaimo, B r i t i s h Columbia; among the hepatics, the only one he reported from B.C. was Marchantia polymorpha L. from Nanaimo. The Catalogue of Canadian Plants (Volume V i i ) published by Macoun (1902) presented a revised l i s t of 57 hepatics from Br i t i s h Columbia, based on Pearson's l i s t and on new collections. Many of the species recorded by Macoun had been sent to L. M. Underwood for identification, and Underwood (1891) had included i n his own l i s t of Hepaticae from the Pacific Coast the records of Macoun's plants with many other collections made to that time. The area encompassed i n this l i s t included several states i n the U.S.A., plus B r i t i s h Columbia, for which Underwood l i s t e d 112 species. The next publication dealing with hepatics from B r i t i s h Columbia was an annotated, revised l i s t of 55 species from Vancouver Island (Evans 1906b), which added more recent collections from Port Renfrew made by Gertrude Gibbs, Daisy Hone, and S. A. Skinner. Among the 71 species on the l i s t were several western North American endemics and North Pacific species. About 20 years after the last of these early publications, A. H. Brinkman, having made additional collections, started his series of five significant papers on hepatics of Br i t i s h Columbia. His f i r s t l i s t (Brinkman 1923) was based on Macoun's l i s t , collections in the National 5 Herbarium, Ottawa, Canada, specimens supplied by A. W. Evans, and Brinkman's personal collections. A few years later Brinkman ( l 9 3 l ) published his studies of the genera Metzgeria and Riccardia, including specimens collected in British Columbia. He described a new variety, M. furcata (L.) Dum. var. pacifica (apparently M. temperata Kuw.) with pointed and. gemmiparous thallus apices, from Hastings, British Columbia. Brinkman (193^) expanded his l i s t of hepatics to 25^ species and varieties in a report from British Columbia, Alberta, the Yukon, Alaska, and seven western states of the U.S.A., from California to the Rocky Mountains. The l i s t attempted to indicate relative rarity or commonness and distributions of species. It also attempted to reduce synonomies and incorporate results of recent collections and information from Clark and Frye (1928). Brinkman (1935> 1937) later published two more revisions of the l i s t s of hepatics from the West Coast and British Columbia. Apparently, nothing more regarding the hepatics of British Columbia was published until after 1950• Several brief papers have been published since then, mostly dealing with individual species, e.g., Krajina and Brayshaw (1951), Persson (1958), Schofield (I962), Worley (1969), and, as a result of the research reported here, Godfrey (1976). A few species in the hepatic flora of British Columbia have been studied in detail as parts of modern revisions of certain difficult species groups, i.e., the Frullania tamarisci complex (Hattori 1972), the Plagiochila semidecurrens complex (inoue 1965)1 and Plagiochila major (Nees) Arn., P. rhizophora Hatt., and P. schofieldiana H. Inoue (inoue 1972a). Only two recent papers by W. B. Schofield (1968a, 1968b) have been available as guides to the hepatic flora of British Columbia. Schofield's species-list (1968b), in particular, has been the most complete, modern compilation of species in this province. While i t includes many hepatics from northern British Columbia and from the Queen Charlotte Islands that do not occur in the southwestern part of the province, this l i s t served as the i n i t i a l basis for the present floristic work. A few flor i s t i c studies of Hepaticae, with keys for identification of species, exist for particular areas of the western United States, but were of limited use in this present study. These include Howe's (1899) work on the Hepaticae and Anthocerotae of California, Clark and Frye's (1928) The Liverworts of the Northwest, Sanborn's (1929) The Hepaticae and  Anthocerotae of Western Oregon, and Flowers' (I96I) Hepaticae of Utah. The Clark and Frye (1928) book would seem a solid reference for the present study; however, i t is outdated and often difficult to use. Other isolated papers dealing with hepatics of the western United States are not discussed in detail here. One (Fulford 1936a) deserves mention, however, because i t includes reports of hepatics from Mt. Baker National Forest in northwestern Washington State. Also, Hong (1975) has recently compiled a l i s t of the leafy Hepaticae of Montana, with mention of the occurrence of these species in adjacent states and provinces. Species records from British Columbia mentioned in Hong's paper were apparently obtained from Schofield's (1968b) l i s t and from Clark and Frye (1928), and no specific information regarding locations or habitats i s given. Hepaticological information from regions north of British Columbia is also important to investigations in the flora of this province. The history of such studies in Alaska and the Yukon is well documented by Worley (1970, which includes an updated species-list; 1972) and does not need to be repeated here. Much of the important work on the bryophyte flora of Alaska and the Yukon, which includes discussions of Hepaticae from that region, has been done by H. Persson (1946, 19^9, 1952, 1962a, 7 1962b, 1963, 1968) and Persson and Weber (1958). Other important reports on Alaskan Hepaticae have been prepared by Evans (1914) and Eyerdam (1952). 8 LOCATION OF THE STUDY AREA The area of the present study i s located primarily i n southwestern Bri t i s h Columbia, the westernmost province of Canada, and also includes the adjacent part of northwestern Washington State, U.S.A. Important geographical and topographical place-names within this region are included on the general map of the study area i n Figure 1. Boundaries of the area are essentially the Pacific Ocean on the west; Washington State north of the Skagit River on the south (especially the Mt. Baker region); the western slopes of the Cascade Mountains i n Manning Park to the southeast; the Fraser River Canyon to the east; Cache Creek and Pavilion Lake to the northeast; and Birkenhead Lake and Meager Creek to the north. Within these boundaries are a great diversity of available habitats for hepatics, as well as the greatest density of human population i n B r i t i s h Columbia. The susceptibility of natural habitats to pressures of urbanization and i t s related demands on the environment make i t essential that this area be studied without delay. To provide familiarity with this region and a basis for understanding the hepatic flora, the climate, physiography, geology, and dominant vegetation of the study area are described i n the following sections. Regional climate (macroclimate), physiography, and geology influence the types of vascular-plant communities which develop i n a region. A l l four elements determine the microenvironments available to bryophytes, including Hepati cae. The study area l i e s i n close proximity to the Pacific Ocean, which has a moderating effect on the climate of the adjacent land mass, and from which winds bring abundant moisture onto the land. Lack of temperature Figure 1. Reference map 1 Alta Lake 2 Anacortes 3 Ashlu Creek 4 Austin Pass 5 Bamfield 6 Bellingham 7 Birkenhead Lake 8 Botanie Valley 9 Brandywine F a l l s 10 Brew Lake 11 Bridal Veil F a l l s 12 Brooks Peninsula 13 Bute Inlet 14 Cameron Lake 15 Campbell River 16 Cape Scott 17 Cheakamus River 18 Chehalis River 19 Chilliwack Lake 20 Chilliwack River 21 Courtenay 22 Cowitchan Lake 23 Cowitchan River 24 Deeks Creek • the study area. 25 Delta Bog 26 Elaho River 27 Englishman River 28 Forbidden Plateau 29 Fraser River 30 Fraser River Canyon 31 Gabriola Island 32 Galiano Island 33 Gold River 34 Haney Research Forest 35 Harrison Lake 36 Harvey Meadows 37 Hell's Gate 38 Hope 39 Howe Sound 40 Indian Arm 41 Jervis Inlet 42 Joffre Creek T r a i l 43 Juan de Fuca Strait 44 Kamloops Lake 45 Kanaka Creek 46 Kennedy Lake 47 Kingcome Inlet 48 Knight Inlet 49 Lillooet 50 Lillooet Lake 51 Lillooet River 52 Lindeman Lake 53 L i t t l e Qualicum F a l l s 54 Long Beach (Pacific Rim Nat. Pk.) 55 Lund 56 Lytton 57 Manning Prov. Pk. 58 Meager Creek 59 Mt. Seymour 60 Murrin Prov. Pk. 61 Nanaimo 62 Nimpkish Lake 63 Nooksack River 64 Nootka Island 65 North Cascades Nat. Pk. 66 Pavilion Lake 67 Pemberton 68 P i t t Lake 69 Port Alberni 70 Port Hardy 71 Port McNeill 72 Port Renfrew 73 Powell River 74 Ross Lake 75 Saltspring Island 76 Sayward 77 Sechelt 78 Skagit River 79 Squamish 80 Squamish River 81 Strait of Georgia 82 Strathcona Prov. Pk. 83 Sumas Mountain 84 Texada Island 85 Thompson River 86 Toba Inlet 87 Tofino 88 Ucluelet 89 UBC Endowment Lands 90 Vedder Crossing 91 Victoria 92 Whistler Mountain 93 Widgeon Creek 94 Yale 95 Yellow Aster Meadows 10 extremes and high atmospheric humidity create a favorable environment for a rich hepatic flora. The basic suitability of southwestern British Columbia for Hepaticae i s further enhanced by i t s varied topography, which provides a great diversity of hepatic habitats at low and'subalpine or alpine elevations, and on wet windward slopes and drier rainshadow areas to the east. Pleistocene glaciation altered the pre-glacial landscape and further isolated the Pacific Northwest of North America from adjacent areas, produced new habitats for plant colonization, and presumably furthered the development of new endemic plant taxa. Finally, the dominant vascular plant communities help to create a variety of microenvironments for hepatics. 11 CLIMATE OF THE STUDY AREA Southwestern B r i t i s h Columbia encompasses two climatic regions, which may be regarded as Pacific, lying west of the crest of the Coast and Cascade Mountains, and as Cordilleran, to the east (Hare and Thomas 197*0 • The Pacific climate i s dominated by the onshore flow of westerly Pacific airstreams, and i s characterized by moderate year-round temperatures, lack of temperature extremes, high amounts of r a i n f a l l , especially i n winter, and frequent cloud-cover and fog. West-facing mountain slopes receive the most precipitation, which accumulates as deep snow i n winter. East-facing slopes receive less precipitation. The driest area i n southwestern Br i t i s h Columbia and adjacent Washington includes the Gulf Islands, San Juan Islands, vi c i n i t y of Victoria, and parts of the adjacent mainland, a l l of which l i e i n the rainshadow of the Olympic Mountains of Washington. The Cordilleran climate affects the upper Fraser Canyon and Thompson River Valley, and the region northeast of the study area away from the Coast. The climate of this area i s also affected by Pacific disturbances, but continental airstreams from the Great Basin to the southeast, and from the east as Arctic a i r i n cooler months, are frequent and control winter circulation. This part of the study area l i e s i n the rainshadow of the Coast and Cascade Mountains, and i s consequently drier than the region to the west. Desert-like conditions with great temperature extremes occur in the area between Lytton and Kamloops. Annual precipitation and annual accumulated temperature are shown i n Figures 2 and 3. Both maps were sketched following maps compiled by the Division of Geography, UBC (Chapman and Turner 1956). 12 Figure 2. Annual precipitation i n southwestern B r i t i s h Columbia, given i n centimeters. Measurements i n cm converted from measurements i n inches given by Chapman and Turner (195^). LEGEND 12a Figure 3. Accumulated annual temperature i n southwestern B r i t i s h Columbia: Day-Degrees over 6° C. Day-degrees C.. converted from figures given by Chapman and Turner (1956) for day-degrees over 43° F. LEGEND v r L <833 I 1 ' 1 1 •111111111 • ' 11 II 1 1 11 1 11 11 1 ill!!'!!!!!!^!!!:!1 • • 1389-1667 14 PHYSIOGRAPHY AND GEOLOGY OF THE STUDY AREA General Features and History The area of the present study l i e s within the Gordilleran Region, one of the five major physiographic subdivisions recognized for Canada (Holland 1976). The topography of this area i s extremely varied, with sand beaches and rock c l i f f s along the ocean, fiords, valleys, mountains, canyons (especially that of the Fraser River), rivers, and lakes. Holland (1976) divides B r i t i s h Columbia into several physiographic subdivisions, of which four are represented i n the study area (Figure 4 ) . Also, Holland (1976) l i s t s f i v e generalized types of bedrock for southwestern B r i t i s h Columbia (Figure 4 ). Generalized bedrock geology depicted for northwestern Washington State represents a simplification of information given by McKee (1972). The f i r s t of Holland's (1976) physiographic subdivisions i n this area i s the Outer Mountain Area (l) which Includes the Vancouver Island Mountains and the coastal plain. The highest mountain of this area i s M t . Golden Hinde, elevation 2201 m. The Vancouver Island Ranges are composed of folded pre-Cretaceous sedimentary and volcanic rocks, with numerous granitic batholith intrusions. The mountains have resulted from dissection of a Tertiary erosion surface of low r e l i e f (Holland 1976). The erosion that resulted i n formation of the Tertiary surface also supplied the Oligocene and early Miocene sediments which were deposited on the Estevan Coastal Plain of the west coast of the Island. Where the relatively soft Tertiary or Pleistocene and Recent rocks underlie this coastal plain, i t i s f l a t ; where i t i s underlain by harder, more erosion-resistant rocks, 15 Figure 4.. Physiographic subdivisions and generalized geology of southwestern British Columbia. LEGEND Physiographic Subdivisions: 1 Outer Mountain Area 2 Coastal Trough 3 Coast Mountain Area A Coast Mountains B Cascade Mountains 4 Interior Plateau Generalized Geology: "C V /\ Intrusive igneous rock, Late Mesozoic or Tertiary Folded and faulted volcanic and sedimentary rock, chiefly Mesozoic Flat or gently dipping sedimentary rock, Cretaceous and younger Flat-lying lava and some sedimentary rock, Cenozoic 16 surface features include bluffs and protruding hummocks. The Coastal Trough (2) i s the second of Holland's (1976) subdivisions for the area. The Georgia Depression occupies the largest portion of the Coastal Trough here. Two areas of differing submarine topography and different geology l i e on either side of the Georgia Depression. The Georgia Lowland along the eastern side i s underlain by granitic rocks, Jurassic and older, affected by erosion. Also within this area i s the Fraser Lowland, an area of depositional origin extending from the coast at Vancouver and Bellingham (Washington), almost to Hope. The Nanaimo Lowland, encompassing eastern Vancouver Island and the Gulf Islands, i s underlain by Upper Cretaceous sedimentary rocks, sandstone and conglomerate on the ridges, and shales and softer rocks i n the valleys, except on southern Vancouver Island, which i s granitic. Areas of low r e l i e f on northern Vancouver Island l i e within the Nahwitti Lowland of the Hecate Depression, which extends northward to the area of the Queen Charlotte Islands. Most of the Nahwitti Lowland consists of remnants of dissected Tertiary erosion surface, with volcanic rocks of the Vancouver Group. The Suquash Basin, a triangular area west of Port Hardy, however, i s underlain by more easily eroded Upper Cretaceous sedimentary beds. The Coast Mountain Area (3) subdivision (Holland 1976) encompasses two regions, the Coast Mountains and the Cascade Mountains. The Coast Mountains extend from the Fraser River northward more than 1610 km to the Yukon. The highest mountain of the Coast Range i n southwestern B r i t i s h Columbia, Mt. Waddington (elevation 4017 m) l i e s north of the study area. High mountains nearer or within the study area include Mt. Gilbert (3110 m) and Mt. Garibaldi (2679 m)• Bedrock i n the Coast Mountains comprises sedimentary and volcanic rocks (mid-Jurassic and older) with Coast Intrusions, largely of granodiorite and quartz diorite. The Cascade Mountains extend from northern California through Oregon and Washington into southwestern Br i t i s h Columbia. The Cascades are composed of Palaeozoic and Mesozoic sedimentary and volcanic rocks, which have been folded, metamorphosed, and intruded by granitic batholiths (Holland 1976). Rugged peaks, ridges, and cirques give evidence of extensive alpine glaciation; mountain features also include hanging valleys, tarns, and glaciated rock surfaces. A fourth physiographic subdivision (Holland I976) i s termed here the Interior Plateau ( 4 ) . The only part of this region actually included i n the study area encompasses small portions of the Fraser and Thompson Plateaus between Lytton and the Pavilion Lake road. The bedrock i n these areas i s covered with glacial d r i f t . An outline of the generalized geological history of southwestern Br i t i s h Columbia, as discussed by Holland (1976) i s given i n Table I. Pleistocene Geology Most of B r i t i s h Columbia was covered during the Pleistocene by the Cordilleran ice-sheet. Holland (1976) states that glaciation i n an area of mountainous terrain i s considered to follow a cycle. It begins with formation of small cirque glaciers, followed by expansion of these into mountain and valley glaciers, then formation of a mountain ice-cap whose movement i s controlled by underlying topography, and ultimately, at the maximum stage, by a regional ice-sheet'controlled for the most part by climate. As the glaciers recede, the cycle i s more or less reversed, except that alpine glaciers recede and disappear while lobes of the main ice-sheet persist longer i n colder, protected valleys (Mathews 1951) • Holland (1976) states that the distribution of present-day glaciers, e.g., 18 Table I. Simplified outline of geological history i n southwestern British Columbia. Information from Holland (1976); time-scale and divisions from Raven and Axelrod (1974). AGE PERIOD ERA ( 1 0 6 yrs) (Epoch) GEOLOGICAL EVENTS (Recent) Post-glacial u p l i f t of land. o ca. 2.5 QUATERNARY G l a c i a l erosion; deposits of d r i f t , (Pleistocene) outwash; ice-marginal deposits. H O CM 27 R Y Neogene Low-relief surface uplifted again along Island and Coast Mtns.; increased stream erosion; formation of mountainous topography. C E N 0 65 T E R T I A Paleogene Deposition of sediments within coastal trough and on western side of Island Mtns.; reduction of land surface to one of low r e l i e f ; volcanism on southern Vancouver Island. Ancient erosion land surface uplifted, 2 axes of greatest u p l i f t were along Island and Coast Mtns. (separated by an ancestral coastal trough). o H O CRETACEOUS (Upper) Land surface of low r e l i e f ; erosion of mountains led to formation of sediments which were deposited i n marine and brackish water basins. N 110 O to m s 19 between Bella Coola and Garibaldi, indicates to some extent, the major centers for formation of Pleistocene ice. In the vi c i n i t y of the present study area, ice moved from the Coast Mountains, eastward across the Fraser Plateau; southeastward and southward across the Thompson Plateau into Washington State; westward from the mainland across the northern end of Vancouver Island; southward and southeastwcird down the Strait of Georgia and southwestward across the southern end of Vancouver Island; and southward (Puget Lobe) down into Puget Sound, Washington. Holland (1976) refers to evidence indicating there were at least two major advances of the Cordilleran ice-sheet, separated by an interglacial stage. Late Pleistocene events in southwestern B r i t i s h Columbia and northwestern Washington, as discussed by Armstrong et a l . (1965)1 Ryder (1972), and Hansen and Easterbrook (197^), are summarized i n Table II. While the gl a c i a l ice was present, the land surface was depressed. Since disappearance of the ice-sheet, the land surface has risen and essentially regained equilibrium. Volcanism and Alpine Glaciers Volcanic activity has occurred in B r i t i s h Columbia since mid-Tertiary time. Within the study area, several centers of Pleistocene and Recent volcanic activity extend northward from Mt. Garibaldi. This i s also a region of present-day alpine glaciers and ice-fields i n the Coast Mountains, at the heads of the Lillooet, Elaho, and Squamish Rivers. Glaciers i n this area reached their greatest size since the Pleistocene i n the 18th and 19th centuries and are currently receding (Mathews 1951)• Mt. Garibaldi i t s e l f i s a volcano that formed par t i a l l y on 20 Table II. Late Pleistocene history of southwestern B r i t i s h Columbia and northwestern Washington. Information from Armstrong et a l . (l965)> Ryder (1972), and Hansen and Easterbrook (1974). YEARS B.P. POSTGLACIAL 9,000 Indians living.at Yale. SUMAS STADE 10,000 11,000 Valley glacier confined to eastern part of Fraser Lowland, (a lobe of the Cordilleran glacier) o M <t H O o CO EVERS0N INTERSTATE 13,000 Glaciomarine, marine, and related deposits i n coastal lowlands. Sea level fluctuations. Coast Mtns. continuously glaciated. Pollen: pine, (spruce). I CO is o Rapid ice retreat. Deposits of the last major Cordilleran ice i n Puget Lowland, Fraser Lowland, S and E Vancouver Island. 15,000 Ice reached maximum extent ca. 24 km S of Olympia; covered a l l but highest peaks. Evans Creek Stade: formation of large alpine glaciers i n mountains of western Washington adjoining southern Puget Lowland. 25,000 Ice reached north end of Strait of Georgia. Cooler, moister conditions; depression of treeline i n mountains, increased mountain hemlock and spruce pollen; tundra-like lowlands. OLYMPIA INTERSTADE 29,000? Ice absent from lowlands of NW Washington and SW B r i t i s h Columbia. Southern Puget Lowland with shallow lakes, swamps, floodplains, some forest cover. Cool, moist climate. Pollen: pine, grasses, sedges. 21 Pleistocene gl a c i a l ice. Subsequent melting of the ice resulted i n collapse of the original cone (Mathews 1952). Mts. Gayley and Meagher, north of Garibaldi, are piles of volcanic materials modified by erosion (Holland 1976). While volcanism i n the Garibaldi area ceased at about the time of disappearance of the last ice-sheet (Mathews 1951)> explosions i n Recent time occurred near the head of the Lillooet River (Holland 1976). In the Cascade Mountains south of Bri t i s h Columbia are several glaciated, Pleistocene stratovolcanoes, of which Mt. Baker (elevation 3286 m) i s one (McKee 1972). During 19751 this volcano emitted steam and toxic fumes, melting gla c i a l ice. 22 VEGETATION OF THE STUDY AREA The vascular-plant vegetation of an area affects the hepatic f l o r a through the creation of substrates and microhabitats, thus i t i s discussed i n broad terms below. Development of the modern vegetation of the study area since deglaciation has been determined from pollen records and described for the Puget Lowland, i.e., the area from the U.S.-Canadian border southward to Everett (Hansen and Easterbrook 197*0 i the Fraser Valley at Haney (Mathewes 1973)» a n d the lower Fraser River Canyon near Yale (Mathewes and Rouse 1975)• Earliest dates given i n these studies for vegetation, varying with timing of deglaciation i n the particular areas, are 9920 ± 760 years B.P. i n the Puget Lowland, 12,690 ± 190 B.P. at Haney, and 11,140 ± 260 B.P. near Yale. While species composition varies to some extent i n each area, reflecting local climatic differences and other factors, the trends shown in the three areas are similar. In the Fraser Valley and Canyon areas, Pinus contorta dominated the vegetation immediately following deglaciation, with high amounts of Alnus and Pteridium aquilinum, and with Salix and Shepherdia canadensis. Subsequently, Pinus contorta declined, and Picea, Abies, Alnus and Tsuga mertensiana increased. This assemblage dominated the vegetation u n t i l approximately 10,500 B.P., when Pseudotsuga menziesii became dominant. Also present i n this'vegetation were Corylus, Quercus, Artemisia, and Gramineae; Thu.ja and Tsuga remained, though i n lesser amounts, i n the vegetation at this time. After 6600 B.P., several taxa increased i n abundance, e.g., Thu.ja-Chamaecyparis - type, Alnus, Tsuga  heterophylla, Abies, and Betula. Hansen and Easterbrook (197*0 report a similar sequence of vegetation, however, they group Pinus contorta and Pseudotsuga menziesii as co-dominants i n the earliest pollen assemblage. Heusser ( i960) reports a similar sequence of vegetation for the Pacific Northwest, but regards the period of dominance by Pseudotsuga menziesii along with the presence of Quercus and other deciduous tree species as a period of warmer, drier conditions from 850O-3OOO B.P., termed the Hypsithermal. Mathewes (1973) found no evidence for a Hypsithermal interval i n southwestern Br i t i s h Columbia. He interprets the differences i n vegetation composition i n relation to s o i l formation and the different ecological requirements of individual species. The varied present-day topography and related climatic differences allow formation of diverse types of modern vegetation i n southwestern B r i t i s h Columbia and northwestern Washington. The vegetation of the area of this f l o r a f a l l s within seven biogeoclimatic zones as described by Krajina (19^5)• These are shown on the map i n Figure 5> which i s derived and simplified from Krajina's map, "Biogeoclimatic Zones of B r i t i s h Columbia," published by the British Columbia Ecological Reserves Committee. The zones are extended into northern Washington State, based on a comparison of Krajina's map and that given by Franklin and Dyrness (19&9) for the vegetation of Washington and Oregon. The vegetation zones and characteristic vascular plants, and i n some cases, bryophytes, as described i n the literature, are discussed below. Nomenclature used throughout this flora for vascular plants follows Hitchcock and Cronquist (1973); common names (see also Lyons 1952, Franklin and Dyrness 19&9) 3 X 6 g i v e n with the f i r s t mention of each species. Nomenclature for mosses follows Crum et a l . (1973)• 24 Figure 5- Major zones of vegetation i n southwestern B r i t i s h Columbia and northwestern Washington State. Zones derived and simplified from Krajina (1965. and map, "Biogeoclimatic Zones of B r i t i s h Columbia) and Franklin and Dyrness ( I 9 6 9 ) . LEGEND T i l ' I I . Coastal Douglas F i r Zone Y Coastal Western Hemlock Zone V A A A A y * \ Subalpine Mountain Hemlock Zone ; J Alpine Zone and Glaciers Engelmann Spruce - Subalpine F i r Zone WMM Interior Douglas F i r Zone Ponderosa Pine - Bunchgrass Zone 24a 25 Coastal Douglas F i r Zone The lower mainland of southwestern Br i t i s h Columbia and most of the eastern coast of Vancouver Island l i e i n the rainshadow of the Vancouver Island Mountains and the Olympic Mountains of Washington. The drier conditions at low elevations i n these areas result i n formation of a temperate mediterranean coniferous forest, the Coastal Douglas F i r Zone (Krajina 1965)• Franklin and Dyrness (1969) incorporate this type of vegetation (Puget Sound area) into their Tsuga heterophylla Zone. Pseudotsuga menziesii (Douglas f i r ) i s the dominant tree of this zone. Other characteristic species include Tsuga heterophylla (western hemlock), Abies grandis (grand f i r ) , Cornus n u t t a l l i i (Pacific dogwood), Alnus rubra (red alder) (successional), Gaultheria shallon (salal), Berberis nervosa (Oregon-grape), Ribes sanguineum (red currant), Vaccinium  parvifolium (red huckleberry), Polystichum muniturn (swordfern), Stokesiella oregana, and Hylocomium splendens. Krajina (1965) divides the Coastal Douglas F i r Zone into two subzones. The wetter subzone i s characterized by Arbutus menziesii (madrone), Pinus contorta (lodgepole pine), P. monticola (western white pine), and Thuja plicata (western red cedar). The drier subzone i s characterized by several species including Quercus garryana (Garry oak), Camassia quamash (camas), P l e c t r i t i s congesta (seablush), and Zigadenus  venenosus (death-camas). This latter combination of species i s typically limited to open rocky knolls on southeastern Vancouver Island and on the Gulf Islands, essentially the distributional range of Quercus garryana i n southwestern B r i t i s h Columbia (Hosie 1969). The Great Delta Bog, a raised (though s t i l l topogenous) peat bog (Biggs and Hebda 1976), occurs on the Fraser River delta just south of 26 Vancouver. The remaining natural vegetation encompasses open heathland and wooded areas. Of the several species of trees present, only Pinus  contorta occurs on open heathland. Shrubs and herbs include typical bog species. Coastal Western Hemlock Zone The Coastal Western Hemlock Zone (Krajina 1965) encompasses temperate marine, coniferous rainforest at lower and intermediate elevations i n B r i t i s h Columbia. In the southwestern part of the province, i t occurs on the west coast and northern end of Vancouver Island. Along the eastern coast of Vancouver Island and on the mainland portion of the study area, which are within the rainshadow of the Olympic Mountains and Vancouver Island Mountains, the zone also occurs above the drier Coastal Douglas F i r Zone, but below the Subalpine Mountain Hemlock Zone. The Coastal Western Hemlock Zone i s recognized by a combination of characteristic plants (Krajina I965)» including Tsuga heterophylla, Thuja plicata, Menziesia  ferruginea (false azalea), Blechnum spicant (deer fern), Hylocomium  splendens, Isothecium stoloniferum, Plagiothecium undulatum, and Rhytidiadelphus loreus. Variations i n the vegetation of the Coastal Western Hemlock Zone are encountered at lower elevations i n Strathcona Park, central Vancouver Island (Kojima 1971). Because of the edaphic conditions i n this area, the vegetation shows close a f f i n i t i e s to that of the wetter subzone of the Coastal Douglas F i r Zone, and Pseudotsuga menziesii i s a dominant species in this area. A very distinctive type of vegetation occurs on the west coast of Vancouver Island. This area i s considered s t i l l a part of the Coastal 2? Western Hemlock Zone by Krajina (19^5)• Franklin and Dyrness ( I969), however, do consider the corresponding coastal vegetation i n Washing-con and Oregon separately, as the Picea sitchensis Zone, which occupies a narrow strip of land generally only a few kilometers wide. This coastal area has the mildest climate of any type of vegetation i n the Northwest (Franklin and Dyrness 1969)1 and i s influenced by frequent cloud-cover and fog, and by mist carried on winds off the ocean. The rainforest vegetation includes Picea sitchensis (Sitka spruce) which forms pure stands on a narrow strip immediately facing the ocean on western Vancouver Island, Tsuga heterophylla, Thuja plicata, Alnus rubra, and a dense layer of Gaultheria shallon. Between this coastal band of Sitka spruce forest on western Vancouver Island, and vegetation more typical of the Coastal Western Hemlock Zone further inland, i s an area of frequent Sphagnum bogs that occupy shallow depressions i n the topography (Wade I965)• Forested parts of these bogs have Pinus contorta ( i n weirdly shaped, "lollipop" form), Chamaecyparis  nootkatensis (yellow cedar), Tsuga heterophylla, occasional Thuja plicata, and Taxus brevi f o l i a (western yew). Shrubs and herbs of open areas include typical bog taxa. These Sphagnum bogs studied by Wade (1965) l i e between Tofino and Ucluelet, and are considered by him to have developed as a result of increasing lack of s o i l aeration, formation of hardpans, and subsequent decline of a previous forest vegetation. The bogs are characterized by low moor (topogenous) conditions with some indication of i n i t i a t i o n of high moor conditions. Farther north on Vancouver Island, on the Brooks Peninsula and on Cape Scott, high moor or raised (ombrogenous) and blanket bogs do occur over extensive areas. The vegetation i s basically similar to that described above. 28 Subalpine Mountain Hemlock Zone The Subalpine Mountain Hemlock Zone i s discussed i n detail by Brooke et a l . (1970) for the Coast Mountains, and for the Garibaldi Park region where conditions are somewhat transitional between coastal and continental. This zone occurs at elevations ranging approximately from 915 to 1680 or 1890 m (3000 to 5500 or 6000 f t according to Brooke) i n the southern part of B r i t i s h Columbia, occurring above the Coastal Western Hemlock Zone and below the Alpine Zone. The Subalpine Mountain Hemlock Zone i s characterized by rugged physiography, with rock outcrops, ridges, talus slopes, narrow steep-sided valleys, fast-flowing mountain streams, fiords, and heavy accumulations of winter snow. Characteristic vascular plants include: Tsuga mertensiana (mountain hemlock), Abies amabilis, Chamaecyparis nootkatensis, Vaccinium membranaceum (thin-leaved blueberry), V. ovalifolium (early blueberry), Sorbus sitchensis (Sitka mountain-ash), Menziesia ferruginea, Rubus pedatus ( t r a i l i n g rubus), Cladothamnus  pyroliflorus (copper-bush), Gaultheria humifusa (alpine wintergreen), Veratrum viride (false hellebore), and Hippurus montana (mountain mare's-tail) (Brooke et a l . 1970). Within the Subalpine Mountain Hemlock Zone, two subzones with readily apparent differences i n continuity of forest cover are distinguished, the Forest Subzone and the Parkland Subzone. Forest Subzone. This i s the lower subzone, occurring at elevations of approximately 915-1070 m. Because of i t s lower elevation, the Forest Subzone has a longer frost-free period, a considerably lighter snow-pack, and a longer snow-free period than does the Parkland Subzone. In the lowest parts of the Forest Subzone, species more characteristic of the 29 Coastal Western Hemlock Zone occur along stream-edge seeps and i n stagnant depressions. At higher elevations the characteristic vegetation of the subalpine forest develops, with Tsuga mertensiana. Abies amabilis. Chamaecyparis nootkatensis, Vaccinium alaskaense (Alaska blueberry), Rhododendron albiflorum (white-flowered rhododendron), Sorbus sitchensis, Streptopus roseus (twisted stalk), Clintonia uniflora (queen's cup), and several other herbs. At highest elevations i n the Garibaldi area and i n more continental subalpine locations, species typical of the Subalpine Engelmann Spruce - Subalpine F i r Zone occur. Parkland Subzone. Above the continuous forest, at elevations of approximately 1070 to 1220 m or more, the vegetation i s broken into a mosaic of forest patches, shrubby areas, and meadows, largely as a result of variations i n topography and the subsequent effect on snow-lie, snow-creep, and duration of the snow-pack. On the tops of rocky knolls or other physiographic projections that have the longest snow-free period (about eight months or less) the vegetation most resembles subalpine forest. With a slightly longer snow duration and with continuous irrigation from snow-melt leading to hygric or hydric conditions, forb and heath types of meadow communities (see below) are formed. In parts of the Parkland where the snow duration i s at least nine months, as on unstable slopes, on surfaces covered with rock fragments, and i n snow basins, simpler plant communities are formed. Subalpine Meadows. Several types of mountain meadows, i.e., areas with few or no trees and dominated by herbs or small shrubs, occur within the area of this study. These meadows have been variously described as subalpine (Kuramoto and Bl i s s 1970), part of subalpine parkland (Brooke et 30 a l . 1970), timber-line (Franklin and Dyrness I969, Eady 1971), or alpine (Brink 1959. Archer 1963). Meadows l i e i n an ecotone between subalpine forest or parkland below, and alpine areas above. Two major types of meadow communities seemed to be most obvious during this hepaticological study: drier flower (forb) meadows and moist heather (heath) meadows. The f l o r i s t i c a l l y r i c h flower meadows occur on moist to drier slopes. Species composition varies with moisture conditions and general location. Herbs include Arnica l a t i f o l i a (broadleaf arnica), Caltha leptosepala (white marsh-marigold), C a s t i l l e j a miniata (scarlet paintbrush), Epilobium angustifolium (fireweed), Erigeron peregrinus (subalpine daisy), Erythronlum montanum (avalanche l i l y ) , Heracleum  lanatum (cow-parsnip), Leptarrhena p y r o l i f o l i a (false saxifrage), Lupinus  l a t i f o l i u s var. subalpinus (broadleaf lupine), Valeriana sitchensis (mountain heliotrope), and Veratrum viride. The vegetation of moist heather meadows i s characterized by Gassiope mertensiana (white mountain heather), Fhyllodoce empetriformis (pink mountain heather), Vaccinium  deliciosum (Cascade blueberry), and Luetkea pectinata (partridgefoot). The heather meadows provide the better hepatic habitats. Variations i n the vegetation of subalpine meadows occur i n wet depressions and along streamlets. Alpine Zone Several studies have been carried out i n the Alpine Zone (Krajina I965) i n Garibaldi Park, which i s transitional between coastal and continental conditions (Brink 1959, 1964; Archer 1963; Fraser 1970), and in a more continental location near Kelowna, B r i t i s h Columbia (Eady 1971)» that are helpful in understanding this zone i n the area of this f l o r a . 31 Franklin and Dryness (1969) discuss timberline and alpine areas i n Washington. The Alpine Zone in southwestern B r i t i s h Columbia and i n Washington State i s not comparable to that of the Rocky Mountains of North America, since habitats capable of supporting extensive areas of alpine tundra vegetation do not exist i n the Pacific Northwest (Franklin and Dyrness I969). This i s a result of the relatively recent deglaciation i n high mountains of this region, e.g., approximately 100-125 years ago i n the Garibaldi area (Brink 1964). Undeveloped or unstable s o i l s (Brink 1964, Fraser 19?0) and snow conditions help to keep plant communities i n a state of flux (Archer I963). The Alpine Zone as i t occurs i n southwestern B r i t i s h Columbia and adjacent northwestern Washington may be considered the region lying above continuous forest, parkland, and meadows, and below glaciers or permanent snowfields. The Alpine Zone i s characterized by depressions and basins with late snow-lie, rubble, boulder-fields, bare rock outcrops, rock walls, rock pavements, exposed summits and ridges with thin s o i l , and krummholz tree forms. In areas with glaciers there are outwash aprons, glacio-fluvial terraces, and moraines. Solifluction, regelation, congeliturbation, and needle ice contribute to unstable s o i l conditions (Brink 1964). Drought becomes important on pervious soi l s lacking a continuous summer water supply (Brink 1964). The factor considered most important (Archer I963) i n the Alpine Zone, controlling s o i l formation, type of vegetation, and plant succession i s snow, i.e., presence, depth, and movement of snow, and duration of snowbeds. The influence of snow may be modified to some extent by the topography. Alpine plant associations are discussed by the authors l i s t e d above. Archer (1963) and Krajina (I965) consider the Phyllodoce-Cassiope (heather) 32 association the zonal type for the Alpine Zone. With a reduction i n snow cover, however, subsequent succession would lead to extension of subalpine forest into heathlands (Archer 19&3) a s i s similarly occurring with the establishment of tree species i n heather meadows of the subalpine forest -heath ecotone (Brink 1959)• Older communities on glacial forelands, especially on moraine ridges with shorter (8 months) snow duration, have conifers reaching tree proportions, and Fraser (1970) feels that succession on glacial forelands at ca. 1460-1525 m i n the Garibaldi area would lead to development of a subalpine parkland sensu Brooke et a l . ( l 9 ? 0 ) . Engelmann Spruce - Subalpine F i r Zone This zone (Krajina I965) ^ n B r i t i s h Columbia encompasses vegetation of the subalpine coniferous forest of interior, or continental locations, where i t replaces the mountain hemlock forest of coastal subalpine elevations. Transitions from the Coastal Mountain Hemlock Zone to this zone occur toward the northern and eastern parts of the present study area, at Garibaldi and Manning Parks. Joffre Creek actually l i e s within the boundary of the interior subalpine zone. The interior subalpine zone differs from the coastal subalpine zone largely i n i t s lower mean annual temperature, decreased amount of total annual precipitation and annual snowfall, and higher probability of summer drought and frozen ground i n winter (Krajina 1965» Fraser 1970). Abies lasiocarpa (alpine f i r ) , Picea engelmanii (Engelmann spruce), and Pinus albicaulis (whitebark pine) are characteristic trees of the interior subalpine vegetation that occur i n the study area. With drier, more continental conditions coastal subalpine trees other than Tsuga  mertensiana, i.e., Abies amabilis and Chamaecyparis nootkatensis, drop out 33 of the vegetation. This interior subalpine zone i s termed the Abies lasiocarpa Zone by Franklin and Dyrness (1969). They feel that the zone i s the representative i n the Pacific Northwest of the Abies lasiocarpa - Picea engelmanii forests that are widespread i n the Rocky Mountains (Daubenmire 1952, Oosting 195&). Interior Douglas F i r Zone The Interior Douglas F i r Zone (Krajina I965) occurs at elevations of 610-1340 m i n southwestern B r i t i s h Columbia i n the rainshadow of the Coast Mountains. It i s encountered i n the area of this f l o r a i n the Fraser River Canyon between Boston Bar and Lytton, i n the v i c i n i t y of Pavilion Lake, and east of Pemberton at Birkenhead Lake. Characteristic species of the zone (Tisdale and McLean 1957« Krajina I965) that occur i n the study area include Pseudotsuga menziesii, Pinus ponderosa (ponderosa pine), P. contorta, Thuja plicata, Populus tremuloides (trembling aspen), Acer  glabrum (Rocky Mountain maple), Amelanchier a l n i f o l i a (shadbush), Artemisia frigida (prairie sagewort), Ceanothus velutinus (greasewood), and Agropyron spicatum (bluebunch wheatgrass). Ponderosa Pine - Bunchgrass Zone This zone (Krajina I965)1 analogous to the Pinus ponderosa Zone of Franklin and Dyrness ( l 9 & 9 ) i i - s represented only by a few collections from the northeastern l i m i t of the study area, between Lytton and Cache Creek. This i s the driest of a l l the vegetation zones i n B r i t i s h Columbia, with much of i t s scanty precipitation f a l l i n g as snow in winter (Franklin and Dyrness I969). Plants characteristic of this zone include Pinus ponderosa, Populus tremuloid.es, Pseudotsuga menziesii, Philadelphus lewisii (mock orange), Artemisia tridentata (sagebrush), Ghrysothamnus nauseosus (rabbit-bush), Agropyron spicatum, and Opuntia f r a g i l i s (prickly-pear cactus) (Krajina 1965; Franklin and Dyrness I969; Brayshaw I965, 1970). 35 METHODS General Methods For the purpose of this f l o r a , collections of as many coastal hepatic taxa as could he found were made from as many types of habitats as possible, taking into consideration accessibility and financial expense. Vancouver Island was included i n the study area primarily for habitats on i t s wet west coast, and on i t s dry eastern coast. Collections from mountain areas were made on the mainland. Fewer collections were made toward the Interior along the eastern and northeastern boundaries of the study area, where distribution of species and diversity of the hepatic fl o r a becomes severely restricted. Whenever a collection was made, ecological information was recorded on a prepared form designed to f i t into a pocket-sized f i e l d notebook (see Appendix E). This information was later transferred to collection notebooks and to labels for herbarium packets. Data recorded included geographical location and type of habitat, substrate (by species name when a tree), relative amount of moisture i n the substrate, relative amount of light f a l l i n g on the plants, compass-direction facing (where applicable), and relative abundance of the species at the sit e . Each collection was given a separate number i n the f i e l d . In collections later sorted into two or more species i n the laboratory, e.g.-, when rare or less frequent species were found, the letters A , B, etc. were added after the collection number. The date of the collection was always recorded i n order to provide important phenological information on growth of the gametophyte and production of sex organs and sporophytes. 36 A l l hepatics collected were i n i t i a l l y studied while s t i l l fresh and in l i v i n g condition. This necessitated shorter collecting trips and immediate microscopic examination of the plants upon return from the f i e l d . In the event that specimens could not immediately he studied microscopically, their fresh condition was maintained by keeping them damp (in paper packets loosely placed i n large plastic bags) and cool (in shaded sites i n the f i e l d , or i n the refrigerator after return to the laboratory). A l l hepatics collected were immediately identified to genus, and where possible, to species. Other species found mixed i n the packets were identified and recorded. Study of l i v i n g plants prevents confusion and taxonomic d i f f i c u l t i e s that can arise after the plants become dried as a result of any changes i n color or texture of the plant. Such a case was described, for example, by Steere (1975) where the common Lophozia incisa has been misidentified as the genus Fossombronia. Characteristics of oil-bodies are often very useful i n species identification but oil-bodies i n many species disintegrate as or after the plants dry. In such taxa as Calypogeia trichomanis and Lophozia ventricosa var. s i l v i c o l a , oil-body information i s c r i t i c a l to correct identification, and, i n many other hepatics, information obtained on oil-bodies i s very convenient to use i n species identification. In those hepatics which lack oil-bodies i n fresh condition, this fact must be observed before the plants are dried or one can never be certain whether or not oil-bodies were ever present. Taxa i n which oil-bodies are absent are relatively few and w i l l soon become familiar; however, discovery of the lack of oil-bodies i n one hepatic from the region, Schofieldia monticola, gave an immediate clue that led to eventual description of this plant as a new genus (Godfrey 1976). 37 To avoid these difficulties with dried plants, notes were made during the i n i t i a l examination of the specimens to record any important features that might be lost in drying. Oil-bodies were counted, described, measured, and sketched for most of the species collected in the fi r s t field season; subsequently, sketches and notes on oil-bodies were only made for species in which they were known to be important to identification. Labeled specimens of a l l collections were placed in the herbarium of the Department of Botany, University of British Columbia (UBC). Duplicates are retained in my personal herbarium. In addition to the new collections made during this study, approximately 1800 collections of hepatics from the study area already contained in the UBC Herbarium were examined, and information obtained is included in this flora, especially in the maps of local distribution presented in Appendix C. An additional 330 hepatics collected in southwestern British Columbia in the late 19th century by John Macoun, and now housed in the National Herbarium, Ottawa, were also examined and annotated. Description and Measurement In the descriptions of hepatic taxa given later in this flora, certain descriptive terms and measurements are included. Consistency in the manner in which these observations and measurements are made is very important, particularly in regard to the following criteria: 1. Approximate size of the plant. An attempt to equate relative descriptive terms of size with actual measured ranges is given in Appendix F. Measurement of width is often more useful than that of length. The latter may be difficult to determine since i t may not be possible to decide 38 where the base of the shoot i s (shoots may be broken during removal from the tuft of plants, and the lower part of the shoot commonly becomes etiolated and weak near the base). Relative sizes within a genus may be different from those for various species i n different genera; thus some species of Gephaloziella ( a l l minute) may be larger than others. 2 . Color. Color description i s somewhat subjective, but broad, familiar categories are used here. 3. Leaf size. This i s taken as the width at the widest point, or the length, determined along a line perpendicular to the width, extending from the leaf-insertion at i t s base, to the upper margin of the leaf or lobe apex. Long teeth at lobe apices and long-decurrent basal extensions were not included i n the measurements. In complicate-bilobed leaves, e.g., i n Scapania, measurements are often made for the individual lobes, for one lobe compared with the other, and for the length of the keel. The method used to make these measurements for this f l o r a was derived from that used by Schuster (1953) and i s ill u s t r a t e d i n Appendix G. 4. Sinus depth. This i s the measurement from the base of the sinus to the leaf-lobe apex, divided by the length of the leaf. 5. Cells. Cell sizes given i n this f l o r a were compiled from measurements obtained from a l l plants studied of a particular taxon. In each of several leaves examined, length and width were recorded for approximately 10 to 15 c e l l s . Several c r i t e r i a were used i n selecting c e l l s to be measured: the cells occurred i n leaves taken from the mid-stem region of healthy-looking, "typical" plants; they were located i n the middle of the leaf i f "median," or along the margins, or at the base, depending on c e l l location-type; and only those c e l l s that appeared to be "average" after i n i t i a l l y scanning the entire leaf were used. Because 39 of the last criterion, c e l l s that were obviously abnormally small or large were not measured. The f i n a l size of cells i s given i n this f l o r a as a range, with occasional or infrequent measurements i n parentheses. 6. Trigones. Liverwort cells frequently have corner thickenings called trigones. Such c e l l s are termed collenchymatous, as opposed to leptodermous ce l l s that are thin-walled and lack trigones. Trigones vary i n size and shape among the species, and to some extent, within individual species as a result of apparent environmental variation. The basic types of trigones are il l u s t r a t e d i n Appendix G. 7 . Cuticle. The cuticle on leafy liverworts may vary i n texture, and i s categorized i n this f l o r a as smooth, with no markings; papillose, with small bumps; striate, with linear markings formed by elongate ridges; and varrucose, with large, warty bumps (see Appendix G). 8 . Oil-bodies. Several observations must be made on the oil-bodies: presence or absence, and distribution i n cells; number per c e l l ; color (usually colorless); texture (see Appendix G); and shape and size. Shape i s given here as (roughly) spherical, or as longer than wide, termed ellipsoid or ovoid (egg-shaped). Oil-body measurements were made i n median leaf ce l l s , using about five to ten "typical" c e l l s per leaf. Ranges are given for width and length, with infrequent measurements i n parentheses. The methods used to study liverworts i n this f l o r a are largely derived from those used by Schuster (1953? 1966a), and for additional information, the appropriate sections of these references should be consulted. Hepaticological terms commonly used throughout this f l o r a are defined i n the glossary given i n Appendix H. Other useful glossaries are available i n Macvicar (1926) and Flowers (1961). PART II: THE HEPATIC FLORA OF SOUTHWESTERN BRITISH COLUMBIA 41 FORMAT This f l o r a i s based on studies of my recent (1974-1977) collections (ca. 4000 specimens, most of which contained at least two or three species) and on specimens contained i n the University of B r i t i s h Columbia Herbarium (UBC, ca. 1800 specimens), altogether comprising perhaps as many as 10,000 individual species-records. Among the specimens examined are represented 37 families, 64 genera, 166 species, and 2 additional varieties. A species-list i s presented i n Appendix A. Only those specimens personally examined are reported here, i n order to avoid potential confusion from any misidentification or vague citation of distribution i n the older literature. The order of presentation of the hepatic f l o r a i s as follows: 1. Keys to the Hepaticae and Anthocerotae (to genera, and, wherever possible, directly to species). These i n i t i a l keys are divided into sections, grouping the genera into categories based on major morphological c r i t e r i a . Use of a taxonomic key i s always somewhat subjective, depending to some extent on the user's visualization of the plant. Thus for d i f f i c u l t taxa and for taxa which could be viewed i n more than one way, the keys presented here have been designed, as much as possible, to lead to the correct endpoints by more than one choice of routes. 2. Presentation of the flora i s by family (subfamily i n the Jungermanniaceae), genus (subgenus where applicable), and then species. Families, and subfamilies, where necessary, are arranged i n taxonomic order, generally following Schuster (1966a). Genera, subgenera, and 42 species axe arranged alphabetically. This i s not always philosophically satisfactory, especially i n the Jungermanniaceae. It was decided that an alphabetical arrangement for genera and lower taxa would be most practical in alleviating problems of systematic position and i n providing an easily used hepatic manual. The Hepaticae are followed by the Anthocerotae. 3 . Genera. For genera having only one species i n the study area, the species description follows immediately. If more than one species occur, the genus i s followed f i r s t by a brief generic description, then by a key to the species. Where complex generic problems are involved, these are discussed immediately i n the section. 4. Keys to species. An attempt was made to make these keys as simple and direct as possible by using the particular c r i t e r i a that best define each species. In the case of complex species-groups or d i f f i c u l t i e s with anatomical features, choices based on one or two c r i t e r i a simply cannot be relied upon to indicate the correct or more l i k e l y dichotomy. Where necessary, several c r i t e r i a are included to give the user of the key a better chance to make a correct identification. Also, since this key i s designed primarily for the f l o r a of southwestern B r i t i s h Columbia and northwestern Washington, ecological information for the hepatics as they occur i n this part of North America i s occasionally used wherever i t i s considered helpful. 5. Information on the species. Within a genus or subgenus, species are presented i n alphabetical order. Each species description i s as concise as possible, designed to provide only basic facts c r i t i c a l to or helpful i n identification. The length of the description varies with the complexity inherent i n each species or species-group. Following the descriptions i s given an indication of the phytogeography of each species, with i t s worldwide distribution and i t s distribution within the study area. Regional distribution i s also shown of distribution-maps (Appendix C). Ecological information relating to each species, i . e . , habitat, substrate, and relative moisture and light conditions, i s also presented, based on information gathered during this study. Throughout Part II of this f l o r a , additional references are occasionally l i s t e d for various taxa whenever useful information i s available i n bryological literature. Many of these additional references present important keys or illu s t r a t i o n s , information of hi s t o r i c a l interest, or information from regions pertinent to the present f l o r a but not covered i n the general references cited below. System of Classification The system of classification by which hepatic families and subfamilies are arranged i n this f l o r a largely follows that of Schuster (1966a: 38I-3865 also, 1969a, 1974), which i s a revised version of his earlier treatment (Schuster 1958a). The generic and specific concepts, and citation of authorities, follow Schuster wherever possible. In other cases, Muller (1951-1958), Schofield (1968b), Inoue (1974a, 1976), Grolle (1976), and authors of recent monographs have been consulted. Grolle's (1976) very useful paper presents a c r i t i c a l examination of nomenclature for a l l hepatics known from Europe. Abbreviations used i n citation of authorities follow Sayre et a l . (1964). 44 General References In addition to any available useful papers on individual species and genera which w i l l be mentioned in the appropriate sections, the following floras and taxonomic keys are useful and are suggested as general references: Howe (1899), Macvicar (1926), Clark and Frye (1928), Muller. (1951-1958), Arnell (1956), Schuster (1953, 1966a, 1969a, 197*0, Flowers (1961), and Inoue (1974a, I976). Use of Frye and Clark's (1937-1947) work on hepatics of North America was avoided during this present study because i t contains numerous errors and outdated material. Of the above works on hepatics, Schuster's books are particularly helpful, since they include extensive and detailed information on anatomical variation, ecological habits, and species relationships. Muller's volumes are, likewise, essential, since they cover a l l the European hepatic f l o r a . Macvicar's guide to. the B r i t i s h Hepaticae, though published i n 1926, remains a very useful book, with good keys and descriptions of species. Valuable information on the hepatic f l o r a of Japan i s provided by Mizutani and Hattori's (I969) checklist and Inoue's (1974a, 1976) two books. Additional papers dealing with the Japanese hepatic f l o r a w i l l be cited i n the appropriate species sections. 45 KEYS TO THE HEPATICAE AND ANTHOCEROTAE OF SOUTHWESTERN BRITISH COLUMBIA I. Plants with apparent radial symmetry KEY A I. Plants with bilateral symmetry, leafy or thallose - II II. Plants thallose KEY H II. Plants with leaves and stems I l l III. Leaves entire, toothed, or lobed, but not complicate-bilobed, i.e., not with one lobe sharply folded over the other IV III. Leaves complicate-bilobed, i.e., with one lobe folded upon or beneath the other VII IV. Leaves deeply divided, almost to the base KEY B IV. Leaves entire, toothed, or lobed, but not divided nearly to the base V V. Leaf-insertion incubous KEY C V. Leaf-insertion succubous or transverse VI VI. Leaves lobed or toothed KEY D VI. Leaves and leaf margins entire . . . . . KEY E VII. Smaller lobe of leaf folded over the larger postical or ventral lobe; (leaves succubous to transverse) KEY F VII. Smaller lobe of leaf folded under the larger antical or dorsal lobe; (leaves incubous) KEY G 46 KEY A: APPARENT RADIAL SYMMETRY 1. Leaves spi r a l l y arranged, cylindric, i n transverse section morphologically similar to the stem; (plants rare, hyperoceanic) . . . . . TAKAKIA LEPIDOZIOIDES (p. 60) 1. Leaves not cylindric (or i n transverse section, leaves not morphologically similar to stems) 2 2. Leaves unlobed, variously shaped (ovate, triangular, or diamond-shaped); shoots pure green, fleshy, arising from a rhizomatous base, resembling minute Brussels sprouts; infrequent, alpine HAPLOMITRIUM HOOKERI (p. 62) 2. Leaves clearly lobed; shoots not pure green or fleshy, not arising from a rhizomatous base; habitats various 3 3. Leaves divided v i r t u a l l y to the base into uniseriate lobes . . BLEPHAROSTOMA (p. 65) 3. Leaves not divided nearly to the base and/or leaf lobes 2 or more c e l l s wide at the base 4 4. Leaves bilobed 5 4. Leaves 3-5-lobed 7 5. Plants (dark green to blackish and crawling) often appearing mouldy,, becoming covered with a bluish-while crystalline or fibrous substance; (generally alpine) ANTHELIA (p. I76) 5. Plants not appearing mouldy 6 6. Plants 5-25 cm long, golden-green to golden-brown; leaves falcate-secund; leaf lobes with conspicuous vittae; leaf c e l l s with bulging to sinuate trigones . HERBERTA (p. 63) 6. Plants 5-20 mm high, dark olive- to blackish-green; leaves not falcate-secund; leaf lobes lacking vittae; leaf c e l l s thin-walled or ± evenly thick-walled, lacking trigones . . . , HYGROBIELLA LAXIFOLIA (p. 179) 7. Leaves distinctly, symmetrically 4-lobed, the lobes channeled; (underleaves bifid) GHANDONANTHUS (p. 98) 7. Leaves 3-5-lobed, the lobes not channeled 8 47 8. Plants minute, leafy shoots less than 0.5 mm wide; leaves distant, lobes seldom more than 2 cells wide at base, short and r i g i d KURZIA (p. 79) 8. Plants larger, leafy shoots ca. 1.0-2.5 mm wide; leaves imbricate; lobes more than 2 c e l l s wide at base, long-attenuate or with c i l i a t e margins . . PTILIDIUM (p. 67) KEY B: LEAVES DEEPLY DIVIDED (the sinus descending 0.5 or more times the leaf length) 1. Leaves spirally arranged, cylindric, i n transverse section morphologically similar to stems; (plants rare, hyperoceanic) . . . . . TAKAKIA LEPIDOZIOIDES (p. 60) 1. Leaves not cylindric (or i n transverse section, leaves not morphologically similar to stems) 2 2. Leaves divided virtually to the base into uniseriate lobes; (underleaves similar to leaves) BLEPHAROSTOMA (p. 65) 2. Leaves not divided nearly to base and/or leaf lobes 2 or more ce l l s wide at the base 3 3. Leaves bilobed 4 3. Leaves 3 -5 -lobed 5 4. Plants appearing mouldy, becoming covered with a bluish-white crystalline, or fibrous, substance (occasionally dark green or even blackish, crawling); leaves not falcate-secund; generally alpine ANTHELIA (p. 1?6) k. Plants golden-green to golden-brown; leaves falcate-secund; suboceanic, low elevations HERBERTA (p. 63) 5. Leaves deeply (3~)4-lobed, the lobes channeled, r i g i d ; lobe bases armed with teeth or spines, lobe margins entire; underleaves ± as large as the leaves, bilobed with teeth at the base CHANDONANTHUS (p. 98) 5. Leaves 3-5-l°t>ed, not channeled, r i g i d or lax; lobe bases not armed with spines, lobe margins entire or with c i l i a ; underleaves various 6 48 6. Leaves not "hand-like," the lobes lax, f l a c c i d ; leaf c e l l s with bulging or sinuate-confluent trigones . . . . PTILIDIUM (p. 6?) 6. Leaves "hand-like," divided into 3-5 r i g i d lobes; leaf c e l l s evenly thick-walled, lacking trigones . . . 7 7. Plants minute, leafy shoots less than 0.5 mm wide; leaves distant; lobes of leaves seldom more than 2 ce l l s wide at base; underleaves reduced to 2 or 3 short divisions . KURZIA (p. 79) 7. Plants larger than above; leaves contiguous to ± imbricate; lobes of leaves 3 _5 cells or more wide at the base; underleaves similar to leaves i n size and shape . . . LEPIDOZIA (p. 83) KEY G: LEAVES INGUBOUS; ENTIRE, TOOTHED OR SIMPLY LOBED — NOT COMPLIGATE-BILOBED 1. Leaves dist i n c t l y bilobed . . . 2 1. Leaves entire or toothed at apex only 3 2. Leaves "hand-like," divided into 3 _4 r i g i d lobes; leaf cells ± evenly thick-walled, trigones lacking or small . . LEPIDOZIA (p. 83) 2. Leaves not "hand-like," divided into 3 _5 lax or f l a c c i d lobes (lanceolate-attenuate or with c i l i a on the margins); leaf c e l l s with large, bulging to sinuate-confluent trigones . . . . PTILIDIUM (p. 67) 3. Leaves usually 2-3-toothed at the apex; stems with postical f l a g e l l a ; plants firm, opaque BAZZANIA (p. 70) 3. Leaves entire to 2-toothed at apex; stems without postical f l a g e l l a ; plants rather pale, often ± translucent . GALYPOGEIA (p. 84) KEY D: LEAVES SUGCUBOUS OR TRANSVERSE; TOOTHED OR SIMPLY LOBED — NOT GOMPLICATE-BILOBED 1. Leaves and underleaves similar i n size and shape, the plants thus appearing ± radially symmetrical . . . . . 2 1. Underleaves much smaller than leaves, or underleaves absent . . . . 5 49 2. Plants appearing 'bluish-white and mouldy from presence of a white fibrous coating (or occasionally dark green to blackish, crawling) ANTHELIA (p. 1?6) 2. Plants brownish-green or golden, not "mouldy" 3 3. Leaves 4-lcbed . . CHANDONANTHUS (p. 98) 3. Leaves 2-lobed 4 4. Plants golden-green to golden-brown; stem lacking a hyalodermis; leaves vittate; suboceanic HEPJ3ERTA (p. 63) 4. Plants dull olive- to blackish-green; stem with distinct hyalodermis; leaves not vittate; subalpine-descending . . . . . . . HYGROBIELLA IAXIFOLIA (p. 179 ) 5. Leaves unlobed, the margins dentate PLAGIOCHILA (p. 202) 5. Leaves lobed 6 6. Leaves 3 -4-lobed 7 6. Leaves 2-lobed 13 7. Leaf lobes irregular, the margins undulate, occasionally dentate; oil-bodies very numerous, 20-40 or more per cell . . . LOPHOZIA Subgenus MASSULA (p. 117) 7. Leaf lobes well-defined, the 2-4 lobes easily distinguished; oil-bodies 5~20 per cell 8 8. Leaves predominantly 3-lobed on a single shoot 9 8. Leaves predominantly 4-lobed on a single shoot 10 9. Underleaves present (distinct under microscope or hand-lens) . . LOPHOZIA (ORTHOCAULIS) FLOERKEI (p. 122) 9. Underleaves absent TRITOMARIA (p. 125) 10. 10. Underleaves absent . . . . LOPHOZIA (BARBILOPHOZIA) BARBATA (p. 105) Underleaves present (distinct under microscope or hand-lens) . . 11 50 11. Leaf lobes di s t i n c t l y channeled, the base of each lobe armed with teeth or spines; plants appearing ± radially symmetrical . . CHANDONANTHUS (p. 98) 11. Leaf lobes not distinctly channeled, bases of individual lobes not armed with teeth or spines; plants appearing bi l a t e r a l l y symmetrical 12 12. Plants green to golden-green, relatively large, 2-4 mm broad; postical leaf base bearing long c i l i a which consist of elongate ce l l s ; antical leaf base entire . . . . LOPHOZIA Subgenus BARBILOPHOZIA (p. 104) 12.- Plants brownish-green to inky, blackish, quite small, to 1 mm broad; postical leaf base bearing small c i l i a which consist of quadrate cells; antical leaf base of most leaves with a tooth or hook LOPHOZIA (ORTHOGAULIS) QUADRILOBA (p. 123) 13. Leaves succubously inserted 14 13. Leaves transversely inserted . 31 14. Underleaves present, distinct under compound microscope 15 14. Underleaves absent 26 15. Underleaves generally unlobed, lanceolate or ovate, ± entire-margined 16 15. Underleaves clearly bilobed, the margins or bases dentate or c i l i a t e i n some species 21 l 6 . Leaf cells becoming equally thick-walled, lacking trigones; stems with postical stolons; (rare, occurring i n Sphagnum bogs) GLADOPODIELLA FLUITANS (p. I85) 16. Leaf cells thin-walled, either leptodermous or collenchymatous; stems lacking postical stolons 17 17. Leaves shallowly bilobed, the sinus descending only to 0.1 or 0.2 times the leaf length . . . 18 17. Leaves bilobed for more than 0.2 times the leaf length 19 51 18. Sinus of leaf lunate, descending only to 0.1 times the leaf length; oil-bodies consisting of minute spherules; plants pale, translucent; stems slender HARPANTHUS FL0TOVIANUS (p. 199) 18. Sinus of leaf ± acute, descending to 0.2 times the leaf length; oil-bodies homogeneous (becoming segmented), shining; plants opaque; stems stout, fleshy NARDIA JAPONIGA (p. 145) 19. Leaves ± remote on sterile shoots, generally with a tooth at the antical leaf base; cuticle becoming faintly striate; o i l -bodies ca. 12-28 per cell; paroicous . . . . LOPHOZIA (PROTOLOPHOZIA) ELONGATA (p. 124) 19. Leaves generally imbricate, rarely with a tooth on the antical leaf base; cuticle smooth; oil-bodies lacking; dioicous 20 20. Underleaves distinct, large, almost as long as the leaves; leaf cells evenly thick-walled PLEUROGLADA ALBESCENS (p. 186) 20. Underleaves indistinct, irregularly shaped, flaplike, generally concealed by rhizoids; leaf cells thin-walled, leptodermous . . . SCHOFIELDIA M0NTIC0LA (p. 187) 21. Underleaves similar in size and shape to leaves; plants ± radially symmetrical, mosslike . . . HYGROBIELLA LAXIFOLIA (p. I79) 21. Underleaves much smaller than leaves; plants bilaterally symmetrical 22 22. Lobes of underleaves narrowly lanceolate, the margins entire; (plants opaque, bright yellow-green) . . . GEOCALYX GRAVEOLENS (p. 198) 22. Lobes of underleaves lanceolate-subulate, the margins (especially near the base) dentate or ciliate 23 23. Rhizoids restricted to bases of underleaves; cuticle smooth . . . L0PH0C0LEA (p. 200) 23. Rhizoids scattered along ventral stem; cuticle strongly striate 24 24. Leaf insertion subtransverse; leaves with short c i l i a on postical base and/or teeth on antical base LOPHOZIA Subgenus ORTHOCAULIS (p. 121) 24. Leaf insertion very oblique (succubous) to horizontal; leaves lacking c i l i a on postical base and teeth on antical base . . . . 25 52 25. Apices of leaf lobes acute; oil-bodies pale golden to brownish, 2-6 per cell . . . . LOPHOZIA Subgenus LEIOGOLEA (p. 108) 25. Apices of leaf lobes obtuse to rounded; oil-bodies colorless, rather numerous, 20-30 or more per cell . LOPHOZIA (MASSULA) OETUSA (p. 118) 26. Plants very delicate, fragile, pale green; leaves (almost transparent), antically decurrent; rhizoids reddish-purple; sex organs on dorsal stem surface F0SS0MBR0NIA (p. 226) 26. Plants not delicate and fragile; leaves not antically decurrent; rhizoids colorless; antheridia borne in axils of leaves (male bracts), archegonia protected by perianth at apex of main shoot or branch 27 27. Stems with a hyalodermis, the cortical cells "swollen," much larger than the medullary cells; leaf cells lacking oil-bodies . . . . GEPHALOZIA (p. 180) 27. Stems lacking a hyalodermis; leaf cells with or without oil-bodies 28 28. Plants dark green to brownish, becoming fuscous, blackish, appearing "scorched;" leaves distant, the lobe apices obtuse; leaf cells ± evenly thickened GYMNOCOLEA INFIATA (p. 102) 28. Plant color various, but never blackish or "scorched;" leaves imbricate, the lobe apices acute; leaf cells thin-walled, with or without trigones . . 29 29. Leaf cells collenchymatous, trigones present; stems in transverse section appearing differentiated, the ventral medulla consisting of smaller cells, which become brownish and mycorrhizal LOPHOZIA Subgenus LOPHOZIA (p. 110) 29. Leaf cells leptodermous, trigones generally lacking; stem in transverse section undifferentiated, a l l cells ± similar in size and appearance 30 30. Plants opaque, yellowish- or bluish-green; oil-bodies present, minute, generally ± homogeneous, numerous, 20-40 or more per cell LOPHOZIA Subgenus MASSULA (p. 11?) 30. Plants clear, almost transparent green; oil-bodies lacking . . . SCHOFIELDIA MONTICOLA (p. 187) 53 31. Underleaves similar to leaves; plants appearing radially symmetrical, mosslike, dark green or often becoming bluish-white, "mouldy-looking" ANTHELIA (p. 176) 31. Underleaves lacking or not similar to leaves; plants appearing bi l a t e r a l l y symmetrical, not "mouldy" 32 32. Lobes of leaf somewhat unequal, the antical lobe smaller; leaves ± loosely complicate ANASTROPHYLLUM (p. 96) 32. Lobes of leaves equal i n size; leaves not ± complicate 33 33• Leaf c e l l s ± evenly thick-walled, or entirely thin-walled ' (leptodermous), lacking trigones; perianths distinct, extending well beyond bracts 3^ 33• Leaf c e l l s collenchymatous, trigones present and often large to bulging; perianths, when present, included within and attached to bracts; (bracteole lacking) . 37 34. Underleaves large and distinct, almost as long as the leaves . . PLEUROGLADA ALBESCENS (p. 186) 34. Underleaves absent or small 35 35- Plants pale, clear to whitish-green; stems with a hyalodermis; oil-bodies lacking CEPHALOZIA (p. 180) 35• Plants green, golden, golden-brown, red, or blackish; (thread-l i k e ) ; stems lacking a hyalodermis; oil-bodies present . . . . . 36 36. Plants not arising from a rhizomatous base, lacking flagellate branches; underleaves present or absent; subgynoecial branches lacking; bracteole present; perianth 4-5-plicate CEPHALOZIELLA (p. 191) 36. Plants arising from a rhizomatous base with a few flagellate branches; underleaves absent; subgynoecial branches quite common; bracteole absent; perianth antically sulcate, the apex rounded; (uncommon) EREMONOTUS MYRIOCARPUS (p. 100) 37. Plants silvery, light grayish-green to light golden-green; leaves closely imbricate, erect-appressed . . . GYMNOMITRION (p. 148) 37. Plants green, red, chestnut-brown, or blackish; leaves distant to approximate, ± spreading MARSUPELLA (p. 150) 54 KEY E: LEAVES SUCCUBOUS OR TRANSVERSE, ENTIRE (UNLOBED) 1. Plants with postical stolons; underleaves minute, with slime-papillae ODONTOSCHISM (p. 188) 1. Plants lacking postical stolons; underleaves, i f present, lacking slime-papillae . . . . . . . 2 2. Rhizoids originating from rectangular, purple pads located at regular intervals along the ventral stem surface . . . . . . . . GYROTHYRA UNDERWOODIANA (p. 147) 2. Ventral stem surface not with regularly spaced purple pads . . . . 3 3. Underleaves present (distinct, at least under microscope) 4 3. Underleaves absent or very minute 7 4. Median leaf c e l l s large, 45-50 [im or more wide . . . MYLIA (p. 146) 4. Median leaf c e l l s smaller, 36 |om wide or less 5 5. Rhizoids scattered along ventral stem surface . . . . NARDIA (p. 143) 5- Rhizoids restricted.to bases of underleaves 6 6. Leaves a l l entire or emarginate; dioicous; perianths short, barely exceeding stem-calyptrae, situated on short late r a l shoots? plants pale green, transparent, generally on moist to hygrophytic sites GHILOSCYPHUS POLYANTHOS (p. 197) 6. Leaves entire to emarginate on upper portion of stem, bilobed below; paroicous; perianths long, extending well beyond calyptra, terminating long, leafy shoots; plants deep (yellow-) green, generally growing on decaying wood i n the forest . . . . LOPH0C0LEA HETEROPHYLLA (p. 201) 7. Plants very pale green, thin, ± transparent, delicate, fr a g i l e ; leaves thin, undulate, "ru f f l y ; " sex organs scattered along dorsal stem surface, not enclosed i n protective structures; rhizoids purple F0SS0MBR0NIA (p. 226) 7. Plants small to large, but not delicate, fragile, or "ruf f l y ; " antheridia produced i n axils of modified leaves (bracts), archegonia situated at apex of stem, protected by perianth and leafy bracts; rhizoids colorless (or reddish-purple i n two species) 8 55 8. Leaf-insertion dorsally reaching stem midline; antical margin of leaf strongly reflexed, forming a convex antical fold; perianth strongly laterally compressed . . . . PLAGIOGHILA (p. 202) 8. Leaf-insertion not reaching stem midline dorsally; antical margin of leaf not reflexed; perianth tubular or cylindrical, not laterally compressed 9 9. Perianth mouth and bases of female bracts with long teeth or c i l i a ; bracteole large and conspicuous JAME30NIELLA AUTUMNALIS (p. 128) 9. Perianth mouth and bases of female bracts not ciliate; bracteole absent JUNGERMANNIA (p. 129) KEY F: LEAVES COMPLICATE-BILOBED, SMALL LOBE (LOBULE) DORSAL/ANTICAL 1. Leaf lobes lanceolate-pointed; plants dull, glaucous (bluish-green to brownish) DOUINIA OVATA (p. 162) 1. Leaf lobes blunt or rounded, not lanceolate-pointed; plants not dull and glaucous 2 2. Leaf lobes narrow, ligulate, much longer than wide; dorsal and ventral lobes pointing in different directions, the dorsal lobe oriented parallel to the stem, ventral lobe at an angle to the stem; gemmae rounded-cubic to angulate-stellate; perianths ovate, plicate, the mouth narrowed DIPLOPHYLLUM (p. 158) 2. Leaf lobes variously shaped but not ligulate; dorsal and ventral lobes pointing in ± the same direction, both at angles to the stem; gemmae smooth, (spherical to) ovoid-ellipsoid; perianths frontally compressed, the mouth broad, truncate . . . . SCAPANIA (p. I63) KEY G: LEAVES COMPLICATE-BILOBED, SMALL LOBE (LOBULE) VENTRAL/POSTICAL 1. Plants robust, shoots 2-3 mm wide, 10-15 cm or more long, julaceous, brilliant red to purple; trigones coarse-confluent; lobule inflated, hooded in the upper part; hyperoceanic . . . . . PLEUROZIA PURPUREA (p. 213) 1. Plants not as above, never julaceous 2 2. Underleaves present 3 2. Underleaves absent . . . . 4 3. Underleaves clearly bilobed; lobule helmet-shaped, with a very narrow stalklike basal attachment to stem; plants small, 0.5-1.5 mm wide FRULLANIA (p. 219) 3. Underleaves entire (or obscurely notched at apex); lobule oblong, longly attached to the stem; plants medium to robust, 1.5-4.5 mm wide PORELLA (p. 215) 4. Rhizoids arising from ventral stem; leaf cuticle with very large papillae; lobules inflated, with teeth at apex; o i l -bodies several per c e l l COLOLEJEUNEA MACOUNII (p. 225) 4. Rhizoids arising i n a tuft from (postical) lobules of leaves; leaf cuticle smooth; lobules not inflated, the apex entire; oil-bodies l(-2) per c e l l , very large, brownish . . . RADULA (p. 210) KEY H: PLANTS THALLOSE 1. T h a l l i ± orbicular (pale golden-green), the dorsal surface with (or obscured by) bottle-like involucres, each enclosing an archegonium or antheridium; spores remaining united i n permanent tetrads SPHAEROGARPOS TEXANUS (p. 239) 1. T h a l l i elongate, strapshaped, or forming rosettes; sex organs not contained i n bottles on the dorsal surface (bottles i n Blasia pusilla contain numerous gemmae); mature spores separated, not remaining i n permanent tetrads 2 2. Cells of thallus with several small chloroplasts; sporophyte capsules globose or ± cylindrical, with determinate growth . . . .3 2. Cells of thallus each with 1 large chloroplast; sporophyte capsules long, slender filaments, with indeterminate growth . . . 20 3. T h a l l i ± transparent or translucent, ligh t or dark green, OR t h a l l i pale green, with hairs at least on margins; not forming rosettes; a l l rhizoids smooth-walled 4 3. T h a l l i opaque and/or leathery, strap-shaped or forming rosettes; rhizoids, i f present, of two kinds, smooth-walled and pegged 11 57 4. Th a l l i with hairs on margins, midrib, or thallus surface 5 4. T h a l l i lacking hairs along margins or midrib . . . 6 5. Both dorsal and ventral surfaces of thallus densely hairy . . . . APOMETZGERIA PUBESGENS (p. 237) 5. Hairs restricted to margins and midrib of thallus, and mostly on the ventral side METZGERIA (p. 238) 6. Plants very fr a g i l e , delicate, appearing leafy, the t h a l l i very deeply divided; the "leaves" ± transparent, succubous to horizontally inserted; rhizoids purple; archegonia and antheridia exposed, scattered along dorsal surface of stem . . . FOSSOMBRONIA (p. 226) 6. Plants thin or fleshy, d i s t i n c t l y thallose, never so deeply divided as to appear leafy; rhizoids generally colorless or brownish (reddish-purple i n Moerckia b l y t t i i ) ; archegonia and antheridia protected by various structures 7 7. Thallus margins f l a t , regularly scalloped, with rounded lobes, each of which contains a dark bluish-black dot (Nostoc embedded in thallus tissue) BLASIA PUSILLA (p. 228) 7. Thallus margins f l a t to strongly undulate, but not regularly scalloped, lacking bluish-black dots 8 8. Thal l i with abundant rhizoids; gynoecia and androecia on dorsal thallus surface 9 8. Thal l i essentially lacking rhizoids; gynoecia and androecia on short l a t e r a l branches 10 9. Female involucre lobed or laciniate to base, scalelike; antheridia beneath toothed scales on dorsal surface of thallus MOERCKIA (p. 229) 9. Female involucre cylindrical or tubular (complete or incomplete tube); antheridia embedded beneath warty bumps on dorsal thallus surface; (widespread, common) PELLIA (p. 230) 10. T h a l l i large, ca. 5 ro™ o r more wide, l i t t l e branched, bright opaque yellow green, b r i t t l e , greasy-feeling; cells lacking oil-bodies ANEURA PINGUIS (p. 233) 10. Th a l l i smaller, ca. 2 mm wide or less, much branched, bright to dark green (to brownish or blackish); c e l l s with oil-bodies RICCARDIA (p. 234) 58 11. Air-pores on^dofsal surface of thallus visible with hand-lens or naked eye; t h a l l i strap-like, i n mats or patches; capsules of sporophyte borne on stalked receptacles (except i n Targionia  hypophylla 12 11. Air-pores lacking on dorsal surface of thallus (not visible with hand-lens or naked eye); t h a l l i narrow, often forming rosettes; capsules of sporophyte embedded i n thallus tissue . . . 19 12. T h a l l i frequently with gemmae i n cups on dorsal surface, large, 6-11 mm wide, pure to dark green 13 12. T h a l l i lacking gemmae and gemmae-cups, variable i n size and color 14 13. Gemmae-cups lunate; ventral scales i n 2 rows; archegonia on stalked receptacles, antheridia i n sessile discs; occurring i n greenhouses, nurseries, gardens LUNULARIA GRUGIATA (p. 244) 13. Gemmae-cups circular; ventral scales i n 3 rows on each side of midrib; archegonia borne on stalked, "palm-tree"-like receptacles, antheridia embedded i n stalked disc-shaped receptacles MARCHANTIA POLYMORPHA (p. 246) 14. T h a l l i very large, 5-12 mm wide, bright yellow-green, the dorsal thallus surface di s t i n c t l y patterned; air-pores on swollen mounds of colorless cells, appearing as white dots, 1 per pattern-unit; archegonia i n stalked, cone-shaped receptacles, antheridia embedded i n raised pads on dorsal surface of thallus . . GONOGEPHALUM GONICUM (p. 243) 14. T h a l l i smaller, color various; dorsal surface of thallus not dist i n c t l y patterned; air-pores not on swollen mounds of colorless cells; location of sex organs various 15 15. Cells i n dorsal epidermis of thallus with (bulging) trigones . . 16 15. Cells i n dorsal epidermis of thallus lacking trigones 17 16. Air-pores surrounded by 4-5 concentric rings of cel l s ; a i r -chambers in several layers of thallus tissue; ventral scales obliquely lunate, with 2 linear, acute appendages; female involucres stalked REBOULIA HEMISPHERICA (p. 242) 16. Air-pores surrounded by ca. 4 concentric rings of cells; a i r -chambers i n one layer of thallus tissue; ventral scales obliquely triangular, with a broadly awl-shaped appendage; female involucres sessile, situated near apex of thallus, formed by 2 purple-black clamshell-like valves TARGIONIA HYPOPHYLLA (p. 240) 59 17. Air-pores surrounded by 1-J ± concentric rings of cells; pseudoperianth present, appearing as a ring of pendent lanceolate scales attached to lower margin of carpocephalum and surrounding each sporophyte ASTERELLA (p. 241) 17. Air-pores elaborate, not simply surrounded by concentric rings of cel l s ; pseudoperianth lacking 18 18. Lateral margins and apices of t h a l l i appearing "bearded" from projection of large, white ventral scales; archegonia on stalked receptacle arising well behind thrllus apex, the location made apparent by a ring of white scales; antheridia beneath papillae on dorsal thallus surface; air-pores stellate, the radial walls of the bounding c e l l s strongly thickened ATHALAMIA HYALINA (p. 245) 18. Ventral scales not projecting beyond thallus margins; both archegonia and antheridia situated on stalked receptacles, these arising near apices of t h a l l i ; air-pores compound, i n dorsal view with a cross-shaped opening .. . PREISSIA QUADRATA (p. 247) 19. Lobes of t h a l l i 1-3 mm wide, with or without a median groove; t h a l l i lacking large or distinct ventral scales; o i l - c e l l s and air-pores absent i n microscopic surface view; plants floating i n water or growing i n rosettes on moist s o i l . . . . RICCTA (p. 247) 19. Lobes of t h a l l i 5 -10 mm wide; long, pendent, violet scales present on ventral thallus surface; o i l - c e l l s and air-pores visible under compound microscope; plants floating i n water or stranded on muddy shores RIGGIOGARPUS NATANS (p. 25l) 20. T h a l l i forming rosettes; thallus margins strongly undulate-crispate; dorsal surface of thallus with outgrowths of tissue; mature spores black ANTH0GER0S PUNGTATUS (p. 252) 20. Thalli forming patches or rosettes; thallus margins ± f l a t ; dorsal surface of thallus ± smooth; mature spores yellow to brownish . PHAEOGEROS (p. 253) 60 Glass HEPATICAE Gametophyte leafy or thallose. Cells with numerous small chloroplasts and with oil-bodies (in most genera). Stomates and internal mucilage chambers absent. Antheridia exogenous; archegonia flask-shaped, not embedded. Sporophyte with determinate growth, consisting of a spherical to ovoid capsule, lacking a meristematic basal area and stomates. Capsule contents consisting of elaters (hygroscopic, with one or more distinct spiral thickening bands) and spores. Dehiscence usually by sp l i t t i n g along 4 lines into 4 valves; less often irregular or cleistocarpous. Subclass JUNGERMANNIAE Leafy and undifferentiated, or unspecialized, thallose Hepaticae. Order CALOBRYALES Family TAKAKIACEAE TAKAKIA Hatt. et H. Inoue TAKAKIA LEPIDOZIOIDES Hatt. et H. Inoue Plants small, to 1.0-1.5 mm wide, 1 cm high, slender, delicate; bright green; spice-scented when dried; radially symmetrical, consisting of cylindrical to ellip s o i d a l (in transverse section) stems arising from a rhizomatous base and bearing short, spirally arranged leaves which are morphologically similar to the stems. Leaves b i f i d to the base, with mucilage c e l l s present i n the axils. Oil-bodies present, variable i n size, 61 shape, and number; ca. 10 per c e l l , less than or up to 5 P-ni i n diameter (Hattori and Mizutani 1958b). Asexual reproduction presumably by regeneration of new shoots from caducous leaves. Dioicous. Archegonia lateral or apical, resembling leaf-lobes, ± exposed, surrounded by s t y l i (scales) and young leaves. Male plants and sporophytes unknown. Takakia must be the hepatic discovery of this century. Since i t s collection by N. Takaki and i t s original description by Hattori and Inoue (1958), the plant has both created, and partially solved, problems for botanists. Takakia i s considered very ancient and isolated from other plants, including other bryophytes, by several authors. Schuster (1966a: 263) states, In Takakia . . . we have an association of primitive features found i n the Calobryales (lack of dorsiventrality and associated lack of dimorphism i n the enation-like axial outgrowths, or 'leaves'; lack of rhizoids; lack of perianth; lack of restricted archegonium origin; lack of specialized branching patterns) and the lowest chromosome number [n = 4 ] yet recorded i n a land plant. This chromosome number i s identical with those known in some green algae. As a consequence, there i s now more evidence for assuming the origin of the hepatics from plants with erect, symmetric, nonbilateral gametophytes, lacking rhizoids, lacking a perianth, and lacking any major axial differentiation. Distribution: Disjunct Western North America — Eastern Asia; i n the study area, Hyperoceanic. Ecology: Growing on open, north-facing rock walls which are kept constantly dripping wet by water seepage (and cloud or mist) and a covering of the substrate by algal ooze. References: Hattori 1958, Hattori and Inoue 1958, Hattori and Mizutani 1958b, Persson 1958, Tatuno 1958, Inoue 1960a, Proskauer 1962b, Schuster 1966c, Mizutani I967, Sharp and Hattori I967, Hattorl et a l . 1968, Grubb 1970, Schofield 1972a, Hattori et a l . 1973, Hattori et a l . 1974, and Hebant 1975. Family HAPLOMITRIACEAE HAPLOMITRIUM Nees HAPLOMTTRIUM HOOKERI (Smith) Nees Plants minute to small, less than 1 cm high (at least the v i s i b l e , above-ground parts), pure green, usually occurring as scattered plants; fleshy, resembling minute "Brussels sprouts," arising from a colorless, rhizomatous basal region. Leaves transversely inserted, loosely clasping; produced ± equally around the stem, therefore plants appearing radially symmetrical; variously shaped (ovate, triangular, diamond-shaped); with marginal slime-papillae. Cells thin-walled, lacking trigones. Oil-bodies numerous (16-20 to 25-42 per c e l l ) , homogeneous, minute, variable i n size and shape, i n western North America spherical (3«0-3-5 [Am)» triangulate (4-6 |im on a side), fusiform (1.5-2.5 x 4-9 \xm), and lunate (1.5-2.5 x 4-9 (im) (Worley I969: 229). Asexual reproduction lacking. Dioicous. Antheridia very large, orange, on long stalks. Female plants with bracts similar to the leaves; shoot-calyptra present, cylindrical, smooth. Sporophyte capsule cylindrical, the wall unistratose. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (alpine); i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on s o i l (occasionally with some l i t t e r and humus) on moist but exposed alpine sites; often on north-facing slopes. The species may be locally abundant, or occur as scattered plants. References: Evans 1917c; Fulford and D i l l e r I956; Schuster 1966c, 1971a; Yang 1966; Worley 1969; and Grubb 1970. Order JUNGERMAMLALES Family HERBERTAGEAE HERBERTA S. Gray Plants long, slender, variable i n size and length, from 2 to 25 cm long (Schuster 1966a), golden-green to (usually) rich, warm red-gold to golden-brown; typically growing i n luxuriant cushions. Stems simple or sparingly branched; i n transverse section, with a distinct cortex, consisting of 2-4 rows of thick-walled c e l l s ; medullary cells larger i n diameter than cortical c e l l s . Leaves incubously inserted, falcate, postically secund; deeply b i f i d , the lobes narrowly lanceolate, acute to acuminate; slime-papillae occasional on margins of undivided base of leaf. Underleaves very similar to leaves i n size and shape, causing this genus to appear radially symmetrical and moss-like. C e l l walls thick; trigones bulging and sinuate. Median cells elongated, forming a v i t t a . Oil-bodies homogeneous, ca. 4-8 per c e l l . Asexual reproduction lacking. Dioicous. Perianths deeply lobed like leaves. Wall of sporophyte capsule 4-7 c e l l rows thick. References: Evans 1917b; Schuster 1957a; Miller 1962, I965; Proskauer 1962a; Inoue 1974a. 64 i . Plants variable i n size, to 1 mm wide i n late r a l view; leaves 2-lobed, symmetrical; leaf lobes ca. 10-15 c e l l s wide at base; sinus acute to obtuse, descending ca. 0.60-0.75 x "the length of the leaf; undivided portion of leaf below lobes narrow, straiglit-sided; widespread H. ADUNCA 1. Plants robust, 1-2 mm wide i n lateral view; leaves 2(-3)-lobed; asymmetrical, with one lobe usually broader; leaf lobes ca. 15-30 c e l l s wide at base; sinus more broadly rounded and gibbose than i n H. adunca, descending up to 0.5 x the length of the leaf; undivided portion of leaf below lobes broad, the l a t e r a l margins ± bulging to auriculate; hyperoceanic . . H. SAKURAII HEREERTA ADUNGA (Dicks.) S. Gray (See above key and generic description.) Sporophytes occasional i n epiphytic populations. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia (the species s. l a t . ) . The plants from southwestern B r i t i s h Columbia belong to H. adunca subsp. adunca (Dicks.) S. Gray, whose distribution i s North Pacific. In the study area, Suboceanic. Ecology: Growing on organic substrates (decaying trunks, trunks of l i v i n g trees, on humus on the ground, on peaty bog hummocks), or on inorganic substrates (rock outcrops, c l i f f s , boulders) where i t forms and grows on i t s own humus. Substrate mesic to moist (becoming dry on rock), open or shaded. The species i s very abundant on the west coast of Vancouver Island, where i t often forms pure, golden cushions. HERBERTA SAKURAII (Warnst.) Hatt. (See above key and generic description.) Plants of this species were st e r i l e i n southwestern B r i t i s h Columbia. This species was identified recently among Bri t i s h Columbia hepatics by H. Inoue. Herberta sakuraii can be distinguished from H. adunca by i t s 65 more robust size and by characteristics of i t s leaves, especially the much broader, ± auriculate leaf base. Distribution: North Pacific; i n the study area, Hyperoceanic. Ecology: On the Brooks Peninsula, northwestern Vancouver Island, occurring on peaty humus on a hummock, on a slope i n an extensive lodgepole pine blanket bog; also, occurring on humus on the ground i n the cloud-forest on a mountain slope, elevation 5^6 m. At Effingham Inlet, on the north side of Barclay Sound, occurring on a c l i f f . Substrate moist to wet, open to partially shaded. Family BLEPHAROSTOMAGEAE Grolle (1972a) uses the name Pseudolepicoleaceae Fulf. et J. Tayl. for this family. BLEPHAROSTOMA (Dum.) Dum. emend. Lindb. Plants small, delicate, pale yellowish-green. Stems simple or sparingly branched; branches predominantly l a t e r a l , terminal. Leaves transversely inserted, divided almost to the base (to within less than 1 c e l l of the base) into 2-4 uniseriate c i l i a - l i k e segments 6-14 cells long. Cuticle of leaf cells finely striate. Oil-bodies small ± granular to segmented, 2-8 or more per c e l l . Underleaves similar to leaves. Asexual reproduction occasionally present, by spherical, 1-celled, thick-walled, conidia-like gemmae cut off from apices of lobes of leaves and underleaves. Dioicous or monoicous. Perianth long, the mouth with long c i l i a . 1. Leaves divided into 2-3 c i l i a - l i k e lobes, the lobes often once forked B. ARACHNOIDEUM 1. Leaves divided into (3-)4 c i l i a - l i k e lobes, the lobes simple, not forked B. TRICHOPHYLLUM 66 BLEPHAROSTOMA ARACHNOIDEUM Howe Shoots 500-2200 |im wide, 2-10 mm long. Leaves divided into 2-3 laciniae, the laciniae 360-750 jim (6-11 cells) long, often once forked at about the middle or above. Leaf cells 25-50 x 50-112 |_im, 1.5-4.0 x as long as broad. Gemmae frequent, 25 |im i n greatest diameter. The above information i s taken primarily from Howe's (1899) original description of the species. Plants from the Gibbs' collection (Port Renfrew) seemed to be much longer (to ca. 10 mm long) than Howe had indicated (2-5 mm long). Blepharostoma arachnoideum seems to be larger than western plants of B. trichophyllum. Blepharostoma arachnoideum appears to be a rare species and remains poorly known. A second collection of this species was made i n B r i t i s h Columbia very recently by W. B. Schofield (see distribution-map, Appendix C). Schuster (1966a: 745) states that B. t r i chophyllum i s a variable species and that B. arachnoideum may be a mere subspecies. Distribution: Endemic to Western North America; i n the study area, Rare or Restricted, known only from Port Renfrew (collected by Gertrude Gibbs, July 12, 1901) and Raza Island, Georgia Strait (collected by W. B. Schofield, May 25, 1977). Ecology: Growing on s o i l or rock, i n or adjacent to streams. BLEPHAROSTOMA TRICHOPHYLLUM (L.) Dum. Shoots 700-880 \m wide, to 5~9(-12) mm long. Leaves (3-)4-lobed, the laciniae 450-590 |im (9-14 cells) long on upper leaves, simple and not forked. [Female and male bracts, however, may have forked lobes.] Leaf cells 11-15 to 16-18 p.m wide x 30-45 to 40-60 [im long, averaging ca. 3-4 x as long as broad. Gemmae rare, 12-l6(-23) (im i n diameter. [Measurements taken from Schuster (1966a)]. Distribution: Gircumboreal, Bipolar (B. trichophyllum s. l a t . ) ; i n the study area, Widespread (elevation-independent). Ecology: Growing on organic substrates, i.e., decaying wood (logs, stumps) i n the forest or along stream banks; or occasionally on humus and l i t t e r on the ground; less frequently on trees (Sitka spruce, red alder, bigleaf maple). Also on sand, s o i l , humus, or among mosses over inorganic substrates such as on basic (basaltic, limestone, serpentine) outcrops, c l i f f s , and boulders, and on sandy clay on riverbanks. Sites are usually mesic to moist, or becoming dry when on rock; partially shaded to open. The species i s frequently found as threads growing among other bryophytes, or occasionally (especially when on rock) i n pure tufts. References: Arzeni 19^8, Hollensen l$6k. Family PTILIDIACEAE PTILIDIUM Nees Plants growing i n patches or tufts, variously copper-colored, pinnately branched. Leaves almost transversely inserted. Leaves and underleaves deeply divided into 3~5 lobes, the lobe margins c i l i a t e or lobes very elongated, narrowly lanceolate-attenuate. Trigones very large, bulging-sinuate. Oil-bodies numerous, minute, spherical, homogeneous. Dioicous. Perianths large, cylindrical, the mouth c i l i a t e . 1. Leaves divided into long, narrowly lanceolate, "streamer-like" lobes; lobe margins entire or with 1-2 long c i l i a ; (generally epiphytic, at low and subalpine elevations) P. GALIFORNICUM 1. Leaf lobes not long-lanceolate; lobe margins densely c i l i a t e . . . 2 68 2. Largest (dorsal) lobe of leaf broader, 15-23 cells wide at base; width of this lobe greater than the length of the longer marginal c i l i a of leaf; depth of leaf sinus 0.35-0.50(0.6o); plants robust, growing i n deep, luxuriant cushions on rock . . P. GILIARE 2. Largest (dorsal) lobe of leaf narrower, 6-11 ce l l s wide at base; width of this lobe less than the length of i t s longest margined, c i l i a ; depth of leaf sinus (0.50-)0.65-0.80; plants smaller, growing i n mats, epiphytic or on rock . . . P. PULGHERRIMUM PTILIDIUM CALIFORNICUM (Aust.) Und. et Cook Plants medium-sized, shoots less than 1.5 mm wide, golden-green to golden- or coppery-red, growing i n shallow mats. Leaves with 3-4 much elongated, narrowly lanceolate, streamer-like lobes; lobe margins entire or with 1 or 2 isolated c i l i a . Dioicous. Perianths frequent. Sporophytes produced from May through August, October (subalpine). Distribution: North Pacific; i n the study area, Widespread (elevation-independent). Ecology: Usually epiphytic, occurring on several species of trees. Also found on fa l l e n and decaying logs, stumps, bark, or wood. Rarely growing over rock, e.g., mixed with P. pulcherrimum on humus over boulder. Substrates are dry to mesic, usually partially shaded. References: Hattori 1952b. PTILIDIUM CILIARE (L.) Hampe Plants large, shoots 2-3 mm wide, green to reddish-copper to blackish-brown, growing i n deep, luxuriant cushions (ca. 1.5-2.0 cm deep). Leaves concave, lobed; lobe margins densely c i l i a t e ; width of largest (dorsal) lobe at base 14-23 cells and greater than the length of the longest marginal c i l i a of leaf; sinus depth 0.35-0'5°(°'60) x the length of the leaf. 69 Dioicous. Sporophytes not observed. Distribution: Gircumboreal, Bipolar; i n the study area, Dry Northern-Interior. Ecology: Always on rock substrates, growing on i t s own humus or s o i l over boulders on boulder-slopes, occasionally on rock outcrops. Sites are dry to mesic, mostly open. The species i c abundant to very abundant wherever i t occurs. References: Evans 1906a. PTILIDIUM PULGHERRIMUM (G. Web.) Vainio Plants medium-sized, shoots less than 1.5 mm wide, the color variable, yellow- to golden-green to golden-red or brownish-purple-red to dark copper, growing i n shallow mats. Leaves concave, lobed; lobe margins densely c i l i a t e ; width of largest (dorsal) lobe at base (5)6-11(12,15) cells, and less than the length of i t s longest marginal c i l i a ; sinus depth (0.50)0.66-0.77 x the length of the leaf. Dioicous. Perianths frequent, occurring on approximately 2/3 of the collections. Sporophytes found i n June and October. Distribution: Gircumboreal; i n the study area, Relatively Widespread but not oceanic (essentially low-elevation). Ecology: The most varied of the three species of Ptilidium. Epiphytic on several species of trees. Also found on decaying logs, and frequently on rock (with l i t t e r , lichen, and humus over boulders on boulder-slopes; occasionally on outcrops). References: Evans 1906a. 70 Family LEPIDOZIACEAE References: Hattori and Mizutani 1958a. BAZZANIA S. Gray Plants opaque green to golden or brownish. Stems with frequent postical-intercalary branches and f l a g e l l a . Leaves incubously inserted, the apices usually 2-3-dentate. Underleaves large. Oil-bodies homogeneous or becoming segmented. Asexual reproduction, i f present, by caducous leaves and underleaves. Dioicous, usually s t e r i l e . Sexual branches postical-intercalary. It i s apparent from a survey of the literature on Northern Hemisphere species of Bazzania that problems have existed i n certain species as a result of morphological variation and resulting taxonomic confusion. The confusion, which does not yet appear to have been satisfactorily resolved, seems to center around two species, B. denudata and B. tricrenata. Evans (I923e, 1923f), Fulford (1936b), and Schuster (1969a) discuss the taxonomic history involved. Bazzania denudata was originally described from eastern North America (see Schuster 1969a). Important features of this species include frequent caducous leaves, which leave portions of the stem denuded; plane or only slightly convex leaves; green color of the plants; and 6-12 oil-bodies per c e l l . Bazzania denudata subsp. o v i f o l i a of Japan has been said to be olive- to du l l green or somewhat brownish (Hattori and Mizutani 1958a). Schuster (1969a: 91) somewhat hesitantly includes this subspecies as synonomous with B. denudata, and more recently, Inoue (l97^a) has treated the Japanese and North American populations as taxonomically the same plant. Ideally, B. tricrenata d i f f e r s i n i t s lack of caducous leaves, 71 convex to strongly deflexed leaves, brownish pigmentation, and oil-bodies numbering 2-4 per c e l l . Problems seem to arise from the a b i l i t y of each of the two species to vary greatly, and confusion becomes most acute i n plants having both brownish (or golden- to brownish-green) color and some caducous leaves. Macvicar (1926) ut i l i z e d B. triangularis (Lindb.) Pears, to accomodate such plants; however, Muller (1956) and Schuster (1969a) regard this species as a synonym (and a slender form) of B. tricrenata. A similar problem exists i n western North America, with B. ambigua. Bazzania ambigua i s recognized or at least mentioned i n the literature as a Pacific species known only from the Pacific Northwest of North America (e.g., i n Evans 1923f» Fulford 1936b, and Schuster 1969a). As a result of variation within single plants and among plants from different sites, confusion exists between B. ambigua and B. denudata. While recognizing both species, Evans (l923f: 60) stated, "It would probably be safe to regard B. ambigua as the western representative of B. denudata . . . " Fulford (1936b: 422) says about B. ambigua, "The plants are variable both i n size and i n appearance and are, under some conditions, d i f f i c u l t to distinguish from B. denudata. Well developed specimens of the two can readily be separated." Hattori and Mizutani (1958a: 106) comment that B. ambigua seems to be closely related to, and perhaps conspecific with B. denudata. Species of Bazzania are usually sterile and gametophytic features are typically used to separate species. The features by which B. ambigua and B, denudata supposedly can be distinguished are discussed by Evans (l923e, 1923f) and Fulford (1936b), and are as follows: 1. Color. Bazzania ambigua i s said to be pale yellow-green to brownish, vrhile B. denudata i s pale green or d u l l to olive-green and at 72 least i n eastern North America, "never with brownish pigmentation" (Schuster 1969a: 9l)» However, i n comparing the pigmentation of B. ambigua with B. tricrenata which i s strongly brownish, Fulford (1936b: 422) states that "brown pigmentation i s absent or only slightly developed i n B. ambigua." 2. Growth habit. Bazzania ambigua i s said to grow suberect i n crowded tufts, while plants of B. denudata are prostrate or suberect. 3. Lateral branches. These spread obliquely i n B. ambigua, but widely i n B. denudata. 4. Leaves. In B. ambigua leaves are ± deflexed, with the apices mostly 2-toothed. Leaves of B. denudata are plane (or sometimes ± convex) with apices 2-3-dentate. However, Evans (l923f: 60) states that leaves of B. ambigua are "not very different from the leaves of B. denudata," and (Evans 1923e) that leaf apices i n B. denudata are variable with the 3-toothed condition rarely realized. Schuster (1969a: 68) also discusses variation i n dentition of leaf apices. Grolle (1972b) has recently suggested that i t appears that the teeth on the leaf apex i n Bazzania  ambigua are sharply acute, while those of B. denudata are usually bluntly pointed, but this would not seem to be a reliable feature for distinguishing species. 5. Underleaves. Underleaves i n B. ambigua are supposed to have scarcely bulging, entire margins, while those of B. denudata are ± crenulate with some indication of a tooth or lobe on the late r a l margins. However, Evans (l923f: 6l) states, "On the whole the underleaves [of B. ambigual show no very distinctive features when compared with the a l l i e d species." In regard to B. denudata, Evans (1923©: 93) states that the "underleaves are almost as variable as the leaves," and that "while the straight or slightly bulging sides are usually entire . . . vague crenulations or sharper teeth may be present i n rare instances." 73 6. Median leaf c e l l s . Sizes for B. ambigua are given as 20 pjn, a " t r i f l e less" than i n B. denudata (Evans 1923f: 6l), but as 19 x 30 |im by Fulford (1936b). Median leaf cells i n B. denudata are said to be 25 x 27 \xm (Evans 1923e: 93), 31 x 36 \xm (Fulford 1936b: 412), and 25-28 x 27-34 |im (Schuster 1969a: 93). 7. Trigones. Trigones are small, with straight or concave sides i n both species. From the available literature, then, i t seems that there are no real or reliable distinctions that separate B. ambigua from B. denudata. There remains the question of brownish, caducous-leaved plants and their placement i n B. tricrenata. According to Schuster (1969a), B. tricrenata never occurs with caducous leaves i n eastern North America. In some European literature i t has been said that B, tricrenata may occasionally have caducous leaves as, for example, i n B. triangularis. Also, Muller (1956) includes caducous-leaves forms as B. denudata i n his flora, but his description raises some interesting points. F i r s t , he distinguishes B. denudata from B. tricrenata on the basis of the smaller, more delicate form of the f i r s t species, but not on the basis of i t s color (which i s given as "braungrun" i n B. tricrenata). Other features as given i n Muller's description or shown i n his illus t r a t i o n s seem more typical of B. tricrenata: slightly falcate leaves; bidentate leaf apices, with one large and one small tooth; seemingly acute teeth and sinuses (from the il l u s t r a t i o n s ) ; underleaves with 3-*+-lobed apices and a tendency to have small projections of the lateral margins (illustrated); and the 3 to 4 oil-bodies per c e l l , a low number comparable to Schuster's (1969a) 2-4(5) oil-bodies per c e l l i n B. tricrenata. Although Muller (1956) suggests that his B. denudata remained distinct from B. tricrenata i n culture, i t would 74 seem possible that this "B. denudata" could be a form of B. tricrenata or otherwise different from B. denudata of North America. Schuster (1969a: 94) states, i n regard to B. denudata. "The species i s a polymorphous one; i t i s possible that the European plant . . . should be segregated." Grolle (1972b) has recently stated that Bazzania denudata i s lacking from Europe and that records of this species i n central Europe belong mainly to Bazzania flaccida (Dum.) Grolle. According to Schuster (1969a) and Grolle (1972b), caducous-leaved forms of Bazzania tricrenata do not exist i n eastern North America or i n Europe. As a result of this present study, however, there seem to be caducous-leaved Bazzanias known i n southwestern B r i t i s h Columbia, which have the true brownish pigmentation and other features of B. tricrenata (see below). These plants should, i n the future, be compared with the European Bazzania flaccida and studied i n more detail. In the key and descriptions given here, the concept of B. denudata follows that of Schuster (1969a), and that of B. tricrenata largely follows Macvicar (1926), Muller (1956), Hattori and Mizutani (1958a), Schuster (1969a), and Grolle (1972b). Bazzania ambigua i s not recognized i n this flora, but i s considered as confused with B. denudata (and perhaps, to some extent, with B. tricrenata). Bazzania denudata and B. tricrenata show much variation both i n this region and throughout their ranges, and a single, detailed, definitive study of the extent of variation through the entire range of the species would be of great assistance i n further studies of this genus i n the Northern Hemisphere. 1. Plants very robust, 3-6 mm wide B. TRILOBATA 1. Plants small to medium-sized, usually not over 2 mm wide 2 75 2. Antical leaf-base very strongly arched across stem; apices and lateral margins of underleaves entire; trigones very large and confluent, the c e l l cavity stellate B. PEARSONH 2. Antical leaf base not strongly arched across the stem; (or) apices and later a l margins of underleaves usually sinuate to lobulate or dentate; trigones smaller, c e l l cavity rounded . . . . 3 3. Leaves 3-4-dentate at apices; apices and la t e r a l margins of underleaves irregularly lobed or sinuate, appearing ragged . . BAZZANIA (new species) 3. Leaves (l-)2-3~dentate at apices; apices of underleaves subentire, sinuate, or 2-4-lobed; lateral margins entire, sinuate, or with a lobe or tooth, but not ragged 4 4. Leaves plane or slightly convex, ovate, not falcate, frequently caducous, leaving parts of stems denuded; leaf apices with blunt teeth, sinuses shallow and rounded; la t e r a l margins of underleaves usually entire to sinuate; oil-bodies numerous, 6-12 per c e l l ; widespread B. DENUDATA 4. Leaves convex to strongly deflexed, triangular or ovate, ± falcate, usually not (but occasionally) caducous; leaf apices narrowed, oblique, with sharp teeth, the antical tooth larger, sinuses ± acute; lateral margins of underleaves frequently toothed or lobed; oil-bodies fewer, 2-4 per c e l l ; suboceanic . . B. TRICRENATA BAZZANIA DENUDATA (Torr.) Trev. Plants medium-sized, 1-2 mm wide; ligh t pure or yellow-green, or dull golden-green to olive. Branches spreading widely from main stem. Leaves plane to slightly convex, ovate, not falcate; frequently caducous, leaving parts of stems denuded. Leaf apices (l-)2-3-dentate, with blunt teeth, sinuses shallow and rounded. Underleaf apices subentire to sinuate to ± shallowly lobed; l a t e r a l margins usually entire to sinuate. Trigones concave-sided. Oil-bodies homogeneous, numerous, 6-12 per c e l l . Sex organs and sporophytes not observed. Distribution: Disjunct North America — Japan; in the study area, Widespread (essentially low-elevation). 76 Ecology: Common, almost always found in forests, occasionally occurring in bogs or on ocean beaches. Occurring most frequently on organic substrates, especially on decaying or rotting wood, e.g., logs and stumps, also on humus and l i t t e r on the ground and on roots and lower trunks of living trees, especially on red cedar. This species also occurs on inorganic substrates, as on humus and soil over c l i f f s , outcrops, and boulders. The substrate is mesic to moist, usually partially shaded, and the species is usually abundant. References: Muller 1953 (not in Rabenhorst). BAZZANIA PEARSONII Steph. Plants small, 1200-1280 |im wide; light green to golden-green to reddish-brown. Stems 160-240 |im wide. Leaves persistent, with 2-3 sharp teeth at apex; sinus lunate; antical edge of leaf strongly arched across stem. Underleaves quadrate to slightly longer than wide; apex entire and truncate, or with a slight center depression; lateral margins entire to slightly sinuate; width ca. 1.6-2.0 x the width of the stem. Trigones very large, angulate, confluent, the cell cavity becoming stellate. Cuticle smooth to striate. Plants from the study area were sterile. Distribution: Western North America — Western Eurasia — Eastern Asia, (oceanic); in the study area, Hyperoceanic. Ecology: Occurring in a lodgepole pine blanket bog, on peaty humus on hummocks, among Sphagnum and other bryophytes. The substrate was wet with seepage, open to partially shaded. The species was infrequent to abundant where found. 77 BAZZANIA TRICRENATA (Wahlenb.) Lindb. in Brotherus Plants small to medium-sized, 1-2 mm wide, golden- to reddish-brown, or occasionally golden-green to brownish-green. Branches spreading obliquely from main stem; stems l60-240 \m in diameter. Leaves triangular or ovate, ± asymmetrical and falcate, convex to strongly deflexed (the plants then appearing somewhat julaceous); usually persistent, but occasionally becoming caducous (as on small plants at Jones Creek and on the boulder-slope at Bridal Veil Falls). Leaf apices narrowed, oblique, with (l-)2~3 sharp teeth, the antical of which is larger; sinuses tending to be ± acute; antical leaf base arched. Underleaves approximately 2.2-3.5 x as wide as the stem; apices usually with 2-4 narrowly triangular lobes, or occasionally sinuate; lateral margins tending to have lobes or teeth, or occasionally sinuate. Trigones concave-sided or bulging. Oil-bodies homogeneous, 2-4(-6) per cell. Distribution: Circumboreal; in the study area, Suboceanic. Ecology: Occurring in humid locations, usually associated with the forest, otherwise found in bogs. The substrate is organic, and chiefly wood (e.g., decaying logs, roots, and trunk bases of red cedars, or on living Sitka spruce), or peaty bog hummocks with Sphagnum and other mosses, or humusy soil on the ground (cloud-forest on the Brooks Peninsula). The species also grows on humus and among mosses over rock, on c l i f f s , outcrops, or boulders. The substrate is mesic to moist, usually in open (to partially shaded) sites; the species is infrequent to abundant. BAZZANIA TRILOBATA (L.) S. Gray Plants robust, 3 -6 rom wide, green to olive-green, resembling Porella in gross appearance (especially the dry plants). Leaves imbricate, not caducous, hiding the stem, deflexed in dry plants, strongly asymmetric, 78 somewhat falcate j the antical base extending across the stem; postical base becoming ± dilated and auriculate. Leaf apices almost always 3 _ (ientate with sharp teeth; sinuses obtuse to rectangulate. Underleaf apices ragged, irregularly and sharply 4-5-toothed or lobed; l a t e r a l margins ± straight but ragged, sinuate, or coarsely dentate. Trigones large and bulging. Oil-bodies homogeneous, 4-8 per c e l l . Plants from the study area were st e r i l e . Distribution: Circumboreal; i n the study area, Hyperoceanic, known from one collection from northwestern Vancouver Island (R. L. Halbert 5433). Ecology: Growing on partly shaded humus of bog mat, i n an open, mature Sphagnum bog. BAZZANIA (new species) Plants medium-sized, ca. 2000-2160 jim wide; light, pale green. Stems 200-296 |im wide. Leaves persistent, with (2)3(4) teeth at the apex, and with occasional accessory teeth (along the margin or on the apical teeth); sinus broadly U-shaped. Underleaves with 2-4 irregular lobes at the apex, appearing rather ragged; the lateral margins irregularly sinuate to ragged; the width 2 . 0 - 3 . 6 x the width of the stem. Trigones small and concave to slightly bulging. Cuticle smooth. These plants were s t e r i l e . This species w i l l be described i n a future manuscript. Distribution: Endemic to Western North America; i n the study area, Hyperoceanic, known only from the Brooks Peninsula. Ecology: Growing on a decaying log i n a ravine behind an ocean beach, protected by headland c l i f f s . Substrate moist to wet, partially shaded. The species occurred i n abundance. 79 KURZIA von Martens Plants minute to small, the main shoots with leaves ca. I6O-36O |im wide, prostrate to ± ascending, i n pure patches or mixed with other bryophytes; dull-textured, deep or olive-green to brownish. Stems pinnately branched, the lat e r a l branches a l l terminal; also with ventral branches. Main shoots and branches occasionally terminating i n f l a g e l l a . Leaf-insertion transverse to succubous. Leaves handlike, (2-3)4-lobed, the lobes ca. 2-5 c e l l s wide at the base; sinuses descending 0.50-0.95 x the leaf length; lobe apices acute to acuminate. Leaf c e l l s r i g i d , quadrate to oblong, ± thick-walled, lacking trigones. Cuticle faintly papillose to distin c t l y striate. Oil-bodies apparently lacking i n mature leaves (Schuster 1969a). Underleaves transversely inserted; according to Schuster (1969a) varying from 0.20-0.95 x the size of the leaves; (2)3-4-lobed; symmetrical or becoming asymmetric through abortion of 1 or 2 lobes. Asexual reproduction lacking. Dioicous. Sexual branches usually postical. Perianths long, usually fusiform. Sporophytes not observed i n plants from the study area. Much controversy exists i n the literature over the correct name for this genus. Grolle (I963, 1965a, 1976), Mizutani and Hattori (I969), and, recently, Inoue (1974a), use Kurzia von Martens, while other authors, e.g., Evans (1939), Hattori (1952a), Hattori and Mizutani (1958a), Fulford (1966), Schuster (1963, 1969a), and Bonner and Schuster (1964), have used Microlepidozia. The debate over which i s the valid generic name arises partly because Kurzia von Martens was originally (1870) published as a new alga. The concept of Microlepidozia as an entity, f i r s t as a subgenus of Lepidozia, predates its date of publication as a genus i n 1934. Because of this long 80 time of recognition and usage, Bonner and Schuster (1964) and Schuster (1969a) proposed that the name Microlepidozia he conserved. However, as Grolle (1976) states, the proposal to conserve the name Microlepidozia was rejected by the "Committee for Bryophytes" which voted 6 to 2 (with 5 abstentions) i n favor of Kurzia (Taxon 17: 582. I968). Kurzia, therefore, i s the accepted name. For a l l the arguments involved i n the debate over the correct generic name, see Grolle (I963i 1965a), Bonner and Schuster (1964), and Schuster (1969a). While the genus Kurzia i s easy to recognize, identification of the species poses some problems. This i s partly a result of the microscopic nature of distinguishing features, which, i n the available literature, include the shape, number, and form of the lobes of the underleaves, and the degree of dentition or c i l i a production on the perianth-mouth and margins of the female bracts. Only the underleaves were used as c r i t e r i a for species identification i n this study, because female bracts and perianths were not always available, and where they did occur, the extent of c i l i a formation seemed inconclusive. Schuster (1969a) also discusses the variations i n female bracts and perianths of Kurzia (as Microlepidozia) species i n eastern North America, whereby transitional forms occur and morphological distinctions between the two species break down (see especially his Figure 93, P« 56, and p. 62). Apparently, the underleaves are then used for species identification i n such cases. Macvicar (1926) suggests a difference i n the degree of ornamentation of the l e a f - c e l l cuticle, but this feature was not deemed sufficiently reliable to be used for plants from the area of this present study. Schuster (1969a: 57) states, "The entire problem of the variation of M. sylvatica i n the direction of M. trichoclados, M. setacea, and M. makinoana i s examined by Schuster (1968), i n which a key to the four 81 species i s presented." In view of the morphological variation and taxonomic confusion involved i n the genus Kurzia, i t i s unfortunate that this key and discussion were apparently never published. The two species of Kurzia that occur i n eastern North America, K. setacea (Web.) Grolle and K. sylvatica (Evans) Grolle, are ecologically quite different. According to Schuster (1969a: 59)» Kurzia setacea i s "i n North America generally a rather rare species, confined almost wholly to peat bogs." Kurzia sylvatica, however, apparently never occurs i n peat bogs, but grows on peaty s o i l or on peat and Sphagnum over acidic rock (Schuster 1969a). After several reexaminations of specimens from southwestern B r i t i s h Columbia, I have decided that two species occur here, with similar, though often perhaps more subtle, differences i n habitat (see below, under the individual species). The less clear-cut differences i n habitat here perhaps result from the av a i l a b i l i t y of various types of transitional habitats. The two species are separated by the following key: 1. Underleaves of main shoots basically 3 -lobed, but generally with 1 or 2 lobes aborted, i.e., reduced to a length of 1 or 2 cell s and terminated by a slime papilla — therefore, underleaves of main shoots usually consisting of 1 large and 2 much smaller lobes, or of 2 large and 1 much smaller lobe, asymmetric (i.e., the lobes are of different sizes, or i f a l l the same length, 1 or 2 lobes are biseriate, at least toward the base, while the others are uniseriate); [perianth-mouth strongly narrowed, c i l i a short] K. MAKINOANA 1.- Underleaves of main shoots always equally (3~)4-lobed, symmetrical; [perianth-mouth weakly contracted, c i l i a t e -fimbriate with uniseriate c i l i a usually 2-3 ce l l s long; female bracts deeply 3 _4-lobed, the lanceolate lobes acuminate and longly ciliate-fimbriate (Schuster 1969a)] K. SETACEA KURZIA MAKINOANA (Steph.) Grolle (See key and generic description above.) 82 / Hattori and Mizutani (1958a) consider this species to he conspecific with the eastern North American Kurzia sylvatica (Evans) Grolle. Schuster (1969a.) disagrees that the two species are identical, though he admits they share close a f f i n i t y . Inoue (1974a: 177) distinguishes four species of Kurzia i n h i s key. He (pers. comm.) identified the B r i t i s h Columbia plants as K. makinoana. Distribution: North Pacific; i n the study area, Suboceanic. Ecology: Occurring on moist, usually shaded places, i n both forests and bogs, often growing (with or without humus) over decaying wood (logs, stumps, rotting wood on the ground), or on peaty humus i n bog holes or over rock slopes, walls, or rocky pond margins (often with Sphagnum i n such places); infrequently subalpine (Mt. Seymour, Golden Ears, ca. 1430 m). KURZIA SETACEA (Web.) Grolle (See key and generic description above.) Grolle (I963) believes that , on the basis of the apparent habitat of the holotype of this species (Jungermannia setacea G. H. Web., 1778), i.e., a basic, rather than acidic, site, this original plant could not have been Kurzia setacea, but rather must have been K. sylvatica or K. trichoclados. Therefore, he drops the epithet setacea as a "nomen dubium" and accepts the next valid species name, pauciflora (from Jungermannia pauciflora Dicks., 1790), i . e . , i n Kurzia pauciflora (Dicks.) Grolle (Grolle I963, I969, 1976). Schuster (1969a: 59i Footnote 36) examines Grolle 1 s arguments but retains the species epithet setacea because i t has been i n common usage f o r many years. I am using the epithet setacea i n this f l o r a to be consistent with nomenclature used for this species elsewhere i n North America and recently by Inoue (1974a), and, hopefully, to avoid further 83 nomenclatural confusion. Distribution: Disjunct North America — Western Eurasia; i n the study area, S t r i c t l y Oceanic. Ecology: Growing on moist to wet peat, i n open lodgepole pine -Sphagnum bogs and on moist humus on open ground i n cloud-forest on the Brooks Peninsula (elevation ca. 550 m). LEPIDOZIA Dum. emend. Joerg. Plants small to medium-sized, pinnately branched, whitish- or yellowish-green to dark green. Leaves incubously inserted, deeply divided into 3 _5 lobes and resembling l i t t l e "hands." Underleaves similar to leaves. Cells thick-walled; trigones minute or lacking. Oil-bodies small, numerous, ± homogeneous. Asexual reproduction lacking. Dioicous. 1. Plants large, 4-7 mm wide including branches; l a t e r a l branches frequently tapered to f i l i f o r m at apices; leaves almost transversely inserted; leaf lobes broader, 8-14 c e l l s wide at base •. . . L. FTLAMENTOSA 1. Plants smaller, 2-4 mm wide including branches; later a l branches lacking f i l i f o r m ends; leaves obliquely to horizontally inserted; leaf lobes narrower, 4-8 cells wide at the base L. REPTANS LEPIDOZIA FILAMENTOSA (Lehm. et Lindenb.) Gott., Lindenb. et Nees (See key and generic description above.) Plants from the study area were s t e r i l e . Distribution: North Pacific; i n the study area, S t r i c t l y Oceanic. Ecology: On the Brooks Peninsula, found on sand over roots on a high bank over a stream i n the forest; also on humus on the ground i n cloud-forest, elevation 5^ -9 m. The substrate i s mesic to moist, par t i a l l y 84 shaded to open; the species was very abundant where found. LEPIDOZIA REPTANS (L.) Dum. (See key and generic description above.) Sporophytes occasional, March through August at low elevations. Distribution: Gircumboreal; i n the study area, Widespread (elevation-independent). Ecology: Common, generally growing on organic substrates, especially on rotting wood (decaying logs, stumps, well into decay), and on humus and peat. Also on bases of l i v i n g trees. Infrequently found on humus over rock outcrops. Sites are mesic to moist, usually i n the partial shade of forests. Family CALYPOGEIACEAE CALYPOGEIA Raddi emend. Nees Plants ± translucent, pale green. Branching usually ventral-intercalary; ventral stolons and f l a g e l l a lacking. Leaves incubous, apices rounded or shortly bidentate. Leaf c e l l s ± thin-walled, trigones minute to small. Oil-bodies grape-cluster type or segmented. Underleaves relatively large, b i f i d to almost unlobed. Rhizoids restricted to areas at underleaf bases. Gemmae occasionally present, at apices of erect, reduced-leaved shoots. Sex organs borne on ventral-intercalary branches; a long, pendulous, subterranean marsupium developing on female branches. Sporophyte capsules cylindrical, dehiscence by spirally twisting valves. References: Evans 1907a, Inoue 1966, Nolan I969. 85 The genus Galypogeia is, itself, easily distinguished. Difficulties arise, however, in identification of species. Schuster (1969a) discusses the taxonomic confusion involved in this genus, especially in the subgenus Galypogeia, to which species encountered in the region of this flora belong. In spite of much work having been done on Galypogeia, e.g., by Buch (1936, 1942, 1948), Muller (1947, 1956), Bischler (1957)1 and Paton (1962), Schuster (1969a: 124) states, "The taxonomy of the holarctic species of subg. Galypogeia remains subjective." Schuster (1969a: 101, 124-125) points out that Buch and Muller themselves were never able to agree completely on identification and nomenclature of several European species o f Galypogeia. Schuster (1969a: 125) also states that the delimitation of certain taxa in European works did not seem to 'hold' for American phenotypes, and that the European classification (and delimitation of taxa) could not be adopted in toto. Some of the taxonomic problems with the American species of Galypogeia are inherent in the fact that these species appear to be undergoing evolution along lines different from the European populations. Schuster, in his statements quoted above, refers largely or entirely to taxonomic problems arising from comparison of eastern North American species of Galypogeia with the same species from Europe. From a study of the same species from southwestern British Columbia, i t is apparent that one must also consider the type of variation that exists in the plants from western North America. In Table III several major features are compared for the three species which present difficulties in southwestern British Columbia and for the same species from Europe (Muller 1956) and eastern North America (Schuster 1969a). When comparing thickenings on epidermal cells of the capsule wall, the British Columbia plants of Calypogeia muelleriana and C. trichomanis are more similar to the European Table ILL, Comparison of three species of Calypogeia Oil-body color Apices of leaves Underleaves: number of cells separating sinus base from rhizoid-i n i t i a l region Lateral margins of underleaves Underleaf width (ratio) compared with stem width (diameter) Nature of thickenings on epidermal c e l l s of capsule wall i n sporophyte Schuster 1969a colorless acute (to apiculate), usually shortly and narrowly bidentate 1-2 (broad) tooth or angulation 1-5-2.5 alternate long walls ± evenly thick-walled (nodular i n subsp. neogaea of eastern North America) North America and Europe. Calypogeia f i s s a Godfrey (this study) colorless definite tendency to be bidentate or at least pointed 1-3 Muller 1956 colorless acute to bidentate (2 i n illustrations) usual tendency to be dentate or have a "shoulder-like" projection 1.5-2.0 often with a bump or protuberance hardly broader than the stem smooth, sheet-like, on without nodular alternate longer walls thickenings Table IIL (continued) Schuster 1969a Oil-body color colorless Apices of leaves obtusely pointed to narrowly rounded Underleaves: number of ce l l s separating sinus base from rhizoid-i n i t i a l region (3)4-6 Lateral margins of underleaves normally unarmed but (p. 156) "lateral margins frequently angulate or dentate" Underleaf width (ratio) compared with stem width (diameter) 2.2-2.5 Nature of thickenings on epidermal c e l l s of capsule wall i n sporophyte alternate long walls with sheetlike thickenings Calypogeia muelleriana Godfrey (this study) Muller 1956 colorless colorless entire, rounded bluntly acute (3)4-7(8,9) entire occasionally a blunt tooth 2.0-3.5 twice as broad as the stem nodular, on alternate nodular, on alternate long walls long walls TableIIL (continued) Schuster 1969a Oil-body color Apices of leaves Underleaves! number of cel l s separating sinus base from rhizoid-i n i t i a l region Lateral margins of underleaves blue entire, subacute to narrowly rounded, occasionally bidentate 4-5 evenly rounded, rarely with "a broad, vague angulation or obtuse tooth" Underleaf width (ratio) compared with stem width (diameter) Nature of thickenings on epidermal c e l l s of capsule wall i n sporophyte usually 1.5-1.8 (p. 156); 1.8-2.2 (p. 150) nodular thickenings on alternate long walls, with sheet-like extensions of thickening occasionally extending to short walls Galypogeia trichomanis Godfrey (this study) Muller 1956 blue blue usually a few leaves on a plant are ± pointed to emarginate to bidentate bluntly acute, occasionally shortly bidentate (1)2-3 occasionally entire, but without a protuberance a "shoulder-like" projection frequently found on several underleaves of a shoot 1.75-2.00 as broad or somewhat broader smooth (not nodular), without nodular sheet-like, on most (?) thickenings walls — on long walls and on many short walls; not clearly on alternate walls 89 plants than to plants from eastern North America. If underleaves are considered, i t seems than those of B r i t i s h Columbia specimens of C. trichomanis may have a greater frequency of occurrence of a l a t e r a l tooth or protuberance than i n plants from the other regions (thus adding to confusion with C. f i s s a ) . Within the genus Calypogeia, i t seems that the following statements hold true for the species i n southwestern B r i t i s h Columbia. Plants should be identified while s t i l l fresh, or at least the oil-body color should be recorded. (This information, whether the oil-bodies are colorless or blue, should be recorded on the packet label for use i n future hepaticological studies.) Deep blue oil-bodies indicate C. trichomanis and eliminate much confusion. Dead plants lack oil-bodies and may be impossible to identify because separation of C. trichomanis and C. f i s s a i s then d i f f i c u l t or impossible, since both species i n this region appear to have similar tendencies toward acute or bilobed leaf apices and toward protuberances (obtuse teeth) on the l a t e r a l margins of the underleaves. Thickenings on epidermal wall c e l l s of the sporophyte capsule are important taxonomic features, but, unfortunately, sporophytes are not commonly found. In view of my suggestion that C. f i s s a and C. trichomanis may be confused i n the area of this fl o r a , some further statements by Schuster (1969a: I67) should be considered, especially his statement that "Calypogeia  fi s s a , with i t s pattern of variation, therefore, can readily be confused with C. muelleriana. Numerous statements — recent ones included — that there i s intergradation between C. f i s s a and C. 'trichomanis' are based on confusion of the latt e r with C. muelleriana." In this flora, however, a l l determinations of C. trichomanis were made using fresh plants i n which the blue color of the oil-bodies was distinct, and no plants with blue oil-bodies could therefore be confused with species possessing colorless oil-bodies. Schuster (1969a: I67-I68) also states, In Europe, G. f i s s a i s unquestionably much more distinct than i t i s i n the southeastern United States. A l l English specimens of C. f i s s a seen show a remarkably high incidence of bidentate, narrow leaves and have the remote, broad and short, deeply bilobed, l a t e r a l l y dentate underleaves typical of the species. In my experience, such plants are very rare or absent i n North America. Because of morphological differences between G. f i s s a of Europe and of eastern North America, Schuster regards them as being of two different subspecies, i.e., G. f i s s a subsp. f i s s a from Europe, and subsp. neogaea from North America. The B r i t i s h Columbia specimens are more l i k e subsp. fi s s a , based on similarities of the capsule wall. The spores from Br i t i s h Columbia specimens are much smaller than spores from plants from either Europe or eastern North America, i.e., 8-10 i^m i n diameter i n the B. C. plants as compared with 14-16 \m (Europe) and 12.5-14.0 \m (eastern North America) (Schuster 1969a). More sporophytes should be examined i n future collections from B r i t i s h Columbia to check this difference i n size. A comparative study of these three species of Calypogeia from Europe and both eastern and western North America would be extremely valuable. Such a study could be reliable, however, only i f fresh material were used so that oil-body color could be known. Also, sporophytes would have to be studied. Both conditions would be d i f f i c u l t to achieve. Schuster (1969a) refers to a monograph by him which would seem to answer many questions regarding Calypogeia. He states (Schuster 1969a: 125), "A detailed review attempting a complete re-evaluation and introducing new data i s that of Schuster (1968a); i n this there i s a detailed evaluation of taxonomic c r i t e r i a and their relative 'values' and degree of v a r i a b i l i t y . " Unfortunately, i t seems that this monograph did 91 not reach publication. The four species known to occur i n southwestern B r i t i s h Columbia are separated i n the key below. Calypogeia sphagnicola was searched for but was not found i n this area. 1. Oil-bodies deep blue i n fresh plants . C. TRICHOMANIS 1. Oil-bodies colorless (or color unknown) . . 2 2. Underleaves deeply divided into 2 divergent lobes, only 1-3 cell s separating the sinus-base from the r h i z o i d - i n i t i a l region, lateral margins occasionally to usually dentate; apices of leaves obtuse to pointed to bidentate 3 2. Underleaves not so deeply divided, 4 or more cells separating the sinus-base from the r h i z o i d - i n i t i a l region, late r a l margins of underleaf generally entire, rounded; apices of leaves variously rounded, entire 4 3. Plants pale, "smaller and more delicate," (cells) usually translucent; (oil-bodies colorless); gemmae frequent; generally growing on humusy ground i n mesic to hygric sites C. FISSA 3. Plants dark bluish-green, "somewhat larger with larger leaves," (cells) opaque; (oil-bodies deep blue); gemmae infrequent; common on decaying wood and humusy s o i l at low elevations (mesic) and on moist ground i n subalpine meadows . . C. TRICHOMANIS 4. Underleaves virtually entire to emarginate or shallowly bilobed at apex, ca. 10-13 small cells separating sinus-base from the rh i z o i d - i n i t i a l region; growing on humusy s o i l over banks, usually subalpine (?), uncommon C. INTEGRISTIPULA 4. Underleaves distinctly though shallowly bilobed, 4-7 c e l l s separating sinus-base from the r h i z o i d - i n i t i a l region; usually growing on decaying wood at low elevations, rather common C. MUELLERIANA CALYPOGEIA FISSA (L.) Raddi Plants pale, translucent green, 1.5-2.5 mm wide. Leaves variable i n size, with the width ± equal to the length, or slightly broader than long (width equals 1.25-1.30 x the length), or slightly longer than wide (width equals 0.83 x the length). The leaves have a definite tendency to be 92 bidentate at the apex, or at least to be pointed (sometimes obtusely pointed). At least a few leaves on each shoot should show this tendency. Underleaves deeply bilobed to 1-3 c e l l s separating the sinus base from the r h i z o i d - i n i t i a l region, the c e l l s of which occasionally have red pigments i n their walls; underleaves decurrent; the width equals 1.5-2.0 x the width (diameter) of the stem; sinus open, broad; lobes usually widely spreading, with a tendency to be dentate or possess at least a "shoulder-l i k e " projection on the lateral margins. Trigones lacking. Oil-bodies colorless. Gemmae frequent, at tips of attenuated shoots; spherical to ovoid-ellipsoid; 1-2-celled; colorless (green). Sporophytes found i n May. Capsule walls with epidermal cells lacking nodular thickenings; rather the thickenings are smooth, sheet-like, and on alternate longer walls. Elaters b i s p i r a l , 8-10 |im i n diameter. Spores 9-10 u.m i n diameter. Distribution: Disjunct North America — Western Eurasia (the species s. l a t . ) ; the subspecies fissa, to which plants i n southwestern Br i t i s h Columbia apparently belong i s Disjunct Western North America — Western Eurasia; i n the study area, Widespread (elevation-independent). Ecology: Generally growing on moist ground (occasionally over rock) on humus or l i t t e r , or over peat or Sphagnum i n bogs; less frequently growing on decaying wood. Occurring i n habitats where moisture i s plentiful, as on streambanks, i n stream canyons, near waterfalls, over peaty c l i f f ledges, i n bogs, roadside ditches, and occasionally i n forests. Substrate mesic to wet, open to partially shaded; the species i s infrequent to abundant where found. CALYPOGEIA INTEGRISTIPULA Steph. Plants medium to large, ca. 2.0-3.0 mm wide, light green, growing i n 93 patches or mats. Branches postical-intercalary. Leaves imbricate, narrowly ovate, longer than wide; apex narrowly rounded, entire. Underleaves relatively large (when compared with those of other species of Calypogeia i n this fl o r a ) , broadly orbicular, broader than long, the width equal to ca. 2-3 times the width (diameter) of the stem; not decurrent at base; the apex emarginate or shallowly bilobed (in the literature, said to be generally ± entire i n this species); sinus-base separated from prominent r h i z o i d - i n i t i a l region by ca. 10-13 small, ± quadrate-rounded cells (cf. elongate c e l l s i n C. muelleriana); "lobe" apices of underleaves broadly rounded; la t e r a l margins very broadly rounded, entire. Leaf cells thin-walled, lacking trigones. Apical c e l l s occasionally somewhat elongate, forming a part i a l border, 1 c e l l row thick where present, along the leaf margins. Median ce l l s averaging 3^-38 x 38-45 |im [measurements from Schuster (1969a: 204); those from the few collections made from the area of this present study were similar]. Oil-bodies 4-8 per c e l l , grape-cluster type, variable, usually consisting of 8-14 globules, the size variable (Schuster 1969a: 204). Plants from the study area were ster i l e ; no sporophytes were found. Schuster (1969a: 206) states that the species'is autoicous (?) or sometimes paroicous; epidermal c e l l s of capsule wall (at least i n eastern North American plants) with nodular thickenings; elaters ca. 7-8 |im i n diameter; spores 10-12 jam i n diameter. Distribution: Circumboreal5 i n the study area, Subalpine-Descending. Ecology: Growing rather abundantly where found, on s o i l and l i t t e r or humus over banks and on slopes — streambanks, tr a i l s i d e banks i n subalpine forest, and sloping banks i n subalpine meadows, also on a high sand cut-bank over a salt marsh on the Pacific Ocean. Substrate mesic to 94 moist, open to partially shaded. CALYPOGEIA MUELLERIANA (Schiffn.) K. Mull. Plants rather large, 2 mm or more wide; dark green, stems dark. Leaves broadly ovate, often broader than long, with entire, broadly rounded apices. Underleaves quadrate, the width equalling the length to somewha.t broader than long; underleaf width (1.7)2.0-3-5 x the width of the stem; decurrent; bilobed to (3)^-7(8,9) cells separating the sinus base from the rhizoid-initial region; sinus narrowly obtuse; lobes close together, broadly rounded, entire on outer margins, lacking a tendency to be dentate. Median cells variable in size, ca. 26-50 x 5^-80 |im. Trigones lacking or minute. Oil-bodies colorless. Gemmae frequent, on apices of attenuated stem tips, 2-celled, green. Sporophytes found in July. Epidermal cells of capsule wall with nodular thickenings, on alternate long walls. Elaters bispiral, 8-10 |im in diameter. Spores 10-12 \m in diameter. Distribution: Circumboreal (?) (distribution unclear as a result of confusion with other species); in the study area Widespread (essentially low-elevation). Ecology: Generally growing abundantly on decaying logs and stumps in the forest; substrate mesic to moist, in partial shade. Occasionally growing on humus and li t t e r (over soil) on the ground. References: Muller 19^ 0, Arnell 19^8. CALYPOGEIA TRICHOMANIS (L.) Corda Plants variable in size, 1.5-3-5 mm wide; dark bluish-green, with shoot apices occasionally turquoise in fresh material. Leaves broadly ovate-cordate, variable in size, with the length ± equal to the width, or 95 becoming somewhat broader or somewhat longer. Most plants have at least a few leaves that are ± pointed or emarginate. to bidentate at the apex. Underleaves decurrent; ± quadrate, the length equal to the width or to 1.3 x as long as wide; width of underleaf equals (1.5)1.7-2.0(2.l) x the width of the stem; bilobed to (l)2-3 cells separating the sinus base and the r h i z o i d - i n i t i a l area; sinus obtusely rectangular to U-shaped; lobes ± spreading, occasionally broadly rounded and entire on outer margins, but frequently, several underleaves can be found on a plant which have some tendency toward dentition of the later a l margins (this i s usually a "shoulder," l/2 to l/3 the distance down from the apex). Apical cells 20-24(30-34) x 22-40(50) [Am. Median c e l l s of leaf variable, i n two size ranges, 20-32(36) x 28-52 [Am or 30-46 x 46-76 \xm. Trigones lacking or minute. Oil-bodies deep blue, 2-5 per c e l l , segmented or "grape-clusters," ca. 4-5 x 5-10 [Am. Gemmae infrequent, spherical to ovoid, 1-2-celled, green. Sporophytes infrequent, found i n A p r i l and May. Capsule wall with epidermal cells lacking nodular thickenings; thickenings are smooth, sheet-like, and occurring on most (?) of the long walls (not clearly on alternate walls) and on many of the short walls. Elaters b i s p i r a l , 8-10 \xm i n diameter. Spores 10-12 [Am i n diameter. Distribution: Circumboreal (?) (distribution unclear as a result of confusion with other species); i n the study area, Widespread (elevation-independent). Ecology: At low elevations, growing on wood (typically decaying stumps, long), on l i t t e r , or on tree bases, peaty banks i n the forest, streambanks, or i n bogs. Occasionally growing on s o i l , gravel, or sand on the ground or over rock. Such sites are mesic to moist, usually at least 96 partially shaded. At higher elevations, occurring on s o i l and humus on the ground i n open subalpine meadows, on banks of rivulets or lake shores; also occurring on s o i l and humus over rock outcrops i n meadows. These sites are moist to wet, often with seepage or flowing water. References: Evans 1907b, Arnell 1948, Meijer 1953. Family JUNGERMANNIAGEAE It has been customary i n the past, e.g., i n Muller (1954) and Schuster (1969a) to maintain both the Lophoziaceae and Jungermanniaceae as separate families, based largely on lobing of the leaves, tendencies involving development of protective structures for the archegonia and sporophyte, and thickness of the capsule wall of the sporophyte. Transitional situations could be found, however, for a l l features used as family c r i t e r i a . In his revision of the Lophoziaceae of Japan, Kitagawa (1965b) considered Jamesoniella as belonging to Lophoziaceae rather than to Jungermanniaceae. In Volume II of his work on the Hepaticae of eastern North America, Schuster (1969a: 810) pointed out that i n a "conservative" classification scheme, the Lophoziaceae would be reduced i n status to a subfamily, the Lophozioideae, within the family Jungermanniaceae. Schuster (1970) has since discussed and adopted this conservative system, which i s also followed by Grolle (l9?2a, 1976) and Schuster and Damsholt (1974). Subfamily LOPHOZIOIDEAE ANASTROPHYLLUM (Spruce) Steph. Plants small, to 1.5 mm wide, forming loose or stringy mats. Leaf insertion ventrally oblique (succubous), dorsally transverse. Leaves bilobed, concave, vertically oriented. Underleaves lacking (or reduced 97 to slime-papillae). References: Schuster 196la, 1966b. 1. Plants blackish; c e l l s of leaves with confluent, irregular to sinuate trigones and walls; gemmae lacking A. ASSIMILE 1. Plants golden-green to golden- or reddish-brown; cells of leaves usually evenly thick-walled, occasionally with trigones; gemmae usually abundant, golden-red, angular A. MINUTUM ANASTROPHYLLUM ASSIMILE (Mitt.) Steph. » Plants small, ca. 600-1500 |_im wide, dark red-gold to red-brown, forming blackish mats. Stems 12-14 c e l l s high i n transverse section. Leaves concave-conduplicate, obliquely to erect-spreading; the ventral portion obliquely inserted, dorsally transverse and becoming decurrent; lobes asymmetrical, apices incurved. Leaf cells collenchymatous, with sinuate walls and irregular, confluent trigones; elongate at base of leaf, 2-3 x as long as wide. Cuticle smooth. Gemmae lacking. Dioicous. Sporophytes produced i n May on plants at Bridal Veil F a l l s . Seta 9 c e l l s thick. Capsule wall 3 c e l l layers thick. Elaters b i s p i r a l , 6-8 [im i n diameter. Spores 14-15 |im i n diameter. The sporophyte i n this species has only recently been described (Schuster and Damsholt 1974). The data for the B r i t i s h Columbia and Greenland plants vary only slightly, although Schuster and Damsholt (1974: 152) state that the capsule wall i s 4-5-stratose. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — E a s t e r n Asia; i n the study area, Subalpine-Descending. Ecology: Growing on i t s own humus over boulders on a boulder-slope near Bridal Veil Falls; elsewhere, on c l i f f faces. Substrate mesic to 98 moist, open to partially shaded. The species may he locally abundant. ANASTROPHYLLUM MINUTUM (Schreb. ex Cranz) Schust. Plants small, 700-1500 \m wide, golden-green to golden- or reddish-brown. Stems 9-10(ll) c e l l s high. Leaves distant to imbricate, as long as or longer than wide, ± quadrate to short-rectangular, bilobed, concave, ± spreading. Cells predominantly evenly thick-walled, but occasionally with distinct trigones; cuticle papillose. Gemmae abundant, golden-red, ca. 20-26 x 28-36 [am; gemmiparous leaves often becoming ragged-edged. Dioicous. Sporophytes infrequent, September and October (subalpine). Distribution: Circumboreal, Bipolar; i n the study area, Widespread (elevation-independent). Ecology: Growing on rock, i n crevices or on the face of c l i f f s , outcrops, boulders (basaltic, slate, granitic), occasionally with moss humus or some accumulated s o i l ; also, i n bogs, occurring on humus or peat, on the ground or over decaying stumps. The habitats are humid, at low elevations, near lakes, rivers, or streams, or i n bogs, and at subalpine elevations, i n forests or meadows. The substrate i s dry.to mesic, but often subject to seepage, occasionally dripping wet depending on the season; open to partially shaded. CHANDONANTHUS Mitt. Plants f i l i f o r m but r i g i d , forming loose carpets or mats; golden to red-brown. Leaves transversely inserted, deeply 4-lobed, the lobes channeled. Underleaves similar to leaves but b i f i d . Asexual reproduction known only i n C. setiformis (see discussion under that species). Both species are dioicous. 99 References: Schuster i960, Laine 1970. 1. Shoots relatively slender (less than 0.7 mm wide); stem readily visible, not much obscured by leaves; leaf lobes narrowly lanceolate, the middle lobes 8-12 cells wide at the base, apices acute to acuminate, ending in an elongated acute cell; in profile, dorsal surface of basa.l portion of leaf with rigid spines (uniseriate, ciliate spines, terminated by elongate, pointed apical cell, projecting out and back from leaf-lobe bases); leaf cells with confluent trigones, sinuate walls G. FILIFORMIS 1. Shoots relatively robust (more than 0.7 mm wide); stems mostly concealed by leaves; leaf lobes broadly triangular-ovate, the middle lobes 15_30 cells wide at the base, apices obtuse, acute, or rounded, not ending in an elongated, acute cell; in profile, dorsal surface of basal portion of leaf strongly arched, entire or bearing teeth at leaf lobe bases which are broad with 1-4 rows of cells at the base (the apical cell is not differentiated, and teeth do not give the plants a spiny appearance); leaf cells with thin or ± evenly thickened walls, trigones small or corners just rounded. G. SETIFORMIS CHANDONANTHUS FILIFORMIS Steph. (See above key and generic description.) Male plants were found among the specimens from Stawamus Chief, south of Squamish. Distribution: Disjunct Western North America — Eastern Asia; in the study area, Strictly Oceanic. Ecology: Growing on the face and in crevices of rock (including granodiorite) outcrops and c l i f f s . Substrate moist with seepage, in partial to f u l l shade. References: Kitagawa 1973. CHANDONANTHUS SETIFORMIS (Ehrh.) Lindb. (See above key and generic description.) Plants from the study area were sterile. During the course of this study, fasciculate gemmae were discovered 100 on plants of Ghandonanthus setiformis from the Murrin Park area, south of Squamish. This i s the f i r s t known occurrence,of gemmae in the genus, These mature gemmae i n C. setiformis are b r i l l i a n t , deep red to violet-red, l(-2)-celled, angular, varying from triangular to long and spindle-shaped, and 10-26 x 16-44 \im, averaging 16 x 28 |im. In a more or less random count of 50 mature, deep red gemmae from plants collected i n 1974 and 1977, 76$ of the gemmae were 1-celled, while 24$ were 2-celled. The gemmae are formed abundantly on the youngest leaves and underleaves at the shoot apex, i n fascicles and by pinching off of individual c e l l s at leaf lobe apices, along leaf margins, and at tips of leaf c i l i a . Mature leaves which have at one time produced gemmae have much eroded margins and often vary considerably from typical leaves on nongemmiparous plants, i.e., leaves are less deeply divided into lobes, lobe number becomes reduced from 3-4 to 2-3, and there i s no indication of teeth between the lobes at their bases. No gemmae were found on plants of G. f i l i f o r m i s from the area of this f l o r a . Distribution: Gircumboreal - Missing i n Eastern Asia, Arctic-Alpine; i n the study area, Dry Northern-Interior. Ecology: Growing with some accumulated s o i l or humus on boulders on boulder-slopes. Substrate dry to mesic, open or partially shaded under rocks. The species i s locally abundant. EREMONOTUS Lindb. et Kaal. i n Pears. EREMONOTUS MYRIOGARPUS'(Garring.) Lindb. et Kaal. Plants minute, 120-200 p.m wide, ca. 4-10 mm high, (golden-green to) golden-brown. Stems r i g i d , ascending from a rhizomatous base; stem ce l l s 101 ± quadrate, thick-walled. Branching including l a t e r a l "branches, numerous subgynoeciai innovations, and occasional f l a g e l l a . Rhizoids scarce i f present. ~ Leaves transversely inserted, ovate-quadrate, distant and appressed on lower stem, approximate above, divided to below the middle into 2 lanceolate lobes 6-9 cells wide at the base. Median leaf cells at base of leaf lobe even with sinus base ca. 10-14 x 12-16 (im; c e l l walls becoming evenly thickened; cuticle smooth (or occasionally, on some leaves or bracts, strongly papillose). Underleaves absent. Asexual reproduction lacking. Dioicous. Male bracts larger and broader than the leaves, loosely imbricate, the apices mostly obtuse. Female inflorescences terminating main stems and repeatedly innovant branches. Female bracts larger than the leaves, the lobes blunt to acute, margins dentate. Bracteole lacking. Perianth long-exserted, oblong, dorsiventrally compressed with a deep antical sulcus, the apex rounded. Perianth mouth contracted, dentate with long cells, occasionally with teeth consisting of more than one c e l l . Sporophytes not observed. This species was recently determined for the plants from Austin Pass by N. Kitagawa. Distribution: Disjunct Western North America — Western Eurasia — Eastern Asia; i n the study area, S t r i c t l y Subalpine-Alpine. This species i s new to North America. Ecology: Growing on sandy s o i l over basic rock, on outcrops of waterfalls i n subalpine meadows. Substrate mesic to dry, p a r t i a l l y shaded to open. References: Inoue and Kitagawa I96I. 102 GYMN0C0LEA (Dum.) Dum. GYMNOCOLEA INFLATA (Huds.) Dum. Plants medium-sized, 1-2 mm wide, to 3 cm long; dark green to fuscous and blackish; lax. Leaves obliquely (succubous) inserted, f l a t to concave,, distant, bilobed; the lobes obtuse to rounded at apices. Underleaves lacking or very small, reduced. C e l l walls ± thickened, brown; trigones absent or minute; cuticle smooth to striate. Oil-bodies finely granular, ca. 4-8 per c e l l . Gemmae lacking. Dioicous. Perianths frequent, inflated, globose, caducous. Sporophytes not observed. Distribution: Circumboreal - Missing i n Eastern Asia; i n the study area, Widespread (elevation-independent). Ecology: Occurring i n three types of habitats, on wet humus and peat i n pools i n Sphagnum bogs (occasionally over rock); on s o i l or humus i n seeps or pools i n open basaltic rock-slopes and "pavements" (Garibaldi region), and on humus and s o i l i n streamlets, pools, and lake margins (occasionally over rock) i n subalpine meadows. Substrate always wet, usually open. The species i s abundant to very abundant where i t occurs. LOPHOZIA Dum. emend. K. Mull. Plants variable i n size. Shoots occasionally branched, the branching lateral-intercalary and terminal-intercalary; f l a g e l l a or stolons absent. Rhizoids scattered along ventral stem. Leaves succubously inserted, 2-4-lobed. Underleaves present i n some subgenera and species. Asexual reproduction frequent, i n the form of 1-2-celled, usually angulate, gemmae. Dioicous or paroicous. Female bracts larger than leaves; bracteole present. Perianth free from bracts, usually plicate d i s t a l l y ; perigynium 103 lacking. Capsule wall of sporophyte usually 3 _5 _stratose. References: Warnstorf 1919; Evans 1935; Muller 1942a; Schuster 1951; Kitagawa 1962b, 1963b, 1965b, 1966, 196?. Lophozia i s a large genus which, i n i t s broad sense, encompasses a variety of forms. In this present flora, Schuster's (1969a) system of classification i s followed, and Barbilophozia, Orthocaulis, and Leiocolea are retained as subgenera i n Lophozia s. Lat., along with Lophozia, Massula, and Protolophozia. Muller (195^) treated Barbilophozia (including Orthocaulis) independently; Kitagawa (1965b) and Grolle (l9?6) also follow this system. It would be convenient to divide the large genus Lophozia into these several genera, but there are always intermediate, transitional species which seem to negate any ju s t i f i c a t i o n for such subdivision. It i s s t i l l possible to visualize "tendencies" i n the different combinations of features possessed by the various subgenera, and thereby learn to recognize them more quickly. The key below, based on rather idealized concepts and generalized combinations of features, separates the subgenera of Lophozia s. l a t . occurring i n southwestern B r i t i s h Columbia. Even i n this key the problem of delimiting the subgenera i s apparent, as intermediate species do not f i t into any one subgenus. In the treatment that follows the key, subgenera are arranged alphabetically within the genus Lophozia, and species of each subgenus are thus grouped together. 1. Leaves 2-lobed 2 1. Leaves 3-4-lobed 7 2. Underleaves generally absent 3 2. Underleaves present 4 3. Leaf cells collenchymatous; oil-bodies granular, ca. 8-24 per cell; stems, in transverse section differentiated, with ventral cells of the medulla becoming mycorrhizal, brownish . . LOPHOZIA Subg. LOPHOZIA 3. Leaf cells generally leptodermous, lacking trigones; oil-bodies generally homogeneous, 25-40 or more per cell; stems in transverse section undifferentiated . LOPHOZIA Subg. MASSULA 4. Apices of leaf lobes obtuse to rounded 5 4. Apices of leaf lobes ± acute 6 5. Leaves very obliquely to horizontally inserted, ± flat; oil-bodies numerous, 20-30 or more per cell . LOPHOZIA (MASSULA) OBTUSA 5. Leaves slightly succubously to transversely inserted, concave with lobes incurved; oil-bodies 2-4 per cell LOPHOZIA (ORTHOCAULIS) KUNZEANA 6. Leaves frequently with an antical tooth; cuticle of leaf smooth to faintly striate; oil-bodies rather numerous, 12-28 per cell, colorless LOPHOZIA (PROTOLOPHOZIA) ELONGATA 6. Leaves lacking an antical tooth; cuticle of leaf cells very heavily papillose-striate; oil-bodies 2-6 per cell, pale yellowish or golden to brownish LOPHOZIA Subg. LEI0C0LEA 7. Leaf cells generally thin-walled, without trigones; oil-bodies numerous, 25-40 or more per cell LOPHOZIA Subg. MASSULA 7. Leaf cells collenchymatous; oil-bodies mostly 3-8 per cell . . . . 8 8. Leaves nearly horizontally inserted, 4-lobed; c i l i a , i f present on postical leaf base, consisting of narrow, elongate cells LOPHOZIA Subg. BARBILOPHOZIA 8. Leaves subtransversely inserted, (2-)3(-4)-lobed; c i l i a , i f present on postical leaf base, consisting of quadrate cells LOPHOZIA Subg. ORTHOCAULIS LOPHOZIA Subg. BARBILOPHOZIA (Loeske) K. Mull, emend. Buch Plants with 4-lobed leaves, obliquely (succubously) to horizontally inserted; in some species with c i l i a on the postical leaf base, the c i l i a 105 then formed of unlseriate rows of narrow, elongated c e l l s . Underleaves present i n some species. References: Schuster 1951i Grolle i960. Of the three species of Lophozia subg. Barbilophozia occurring i n southwestern B r i t i s h Columbia, L. barbata i s most easily identified (by features given i n the key below). Lophozia hatcheri and L. lycopodioides, however, may be very d i f f i c u l t to separate, and a l l the features included i n the key should be examined, with a decision as to species based on the combination of characters. 1. Leaves ± quadrate, with l a t e r a l margins ± parallel, lobe apices mostly rounded to obtuse; postical leaf base lacking c i l i a ; underleaves lacking or minute and consisting of 1 or 2 filaments of cel l s . L. BARBATA 1. Leaves narrower toward base than i n upper portion, the la t e r a l margins not parallel, lobe apices acute to mucronate; postical leaf base with c i l i a ; underleaves large, lanceolate-ciliate or b i f i d - c i l i a t e 2 2. Plants medium-sized to large, 2-3 mm wide; leaves symmetrical, antical leaf margin ± straight; leaf lobes straight-sided, the middle lobes narrower (width at base averaging ca. 1.2 x the length of the lobe); lobe apices acute to apiculate, the dis t a l c e l l s averaging 4? [im long; postical c i l i a few, usually 2-5» short; gemmae commonly abundant L. HATCHERI 2. Plants robust, ca. 4 mm wide; leaves asymmetrical, antical leaf margin strongly arched; leaf lobes convex-sided, the middle lobes very broad at base (the width averaging 2.0 x the length); lobe apices mucronate, the d i s t a l mucro ce l l s averaging 76 |J.m long; postical c i l i a numerous, frequently 5-8, quite long; gemmae lacking (rare) . . L. LYCOPODIOIDES LOPHOZIA (BARBILOPHOZIA) BARBATA (Schmid.) Dum. Plants large to robust, ca. 3~4 broad; green to golden- or purplish-brown. Stems dark i n appearance, ventral side reddish, with dense rhizoids. Leaves with lateral margins straight, parallel; the base not 106 narrowed; symmetrical-quadrate, flat; normally 4-lobed, occasionally 2-3-lobed, the lobe apices rounded, obtuse, or acute; postical c i l i a lacking. Underleaves lacking or minute and consisting of 1 or 2 cell filaments (or lanceolate segments). Gemmae lacking. Sporophytes found once, in June. Distribution: Gircumboreal; in the study area, Dry Northern-Interior. Ecology: Generally associated with rock; not a pioneer, but occurring on humus, litter, or among mosses, over boulders or outcrops. Infrequently on sandy soil on streambanks; once found on sandy soil and mosses over a decaying log. In forests, on boulder-slopes, where the substrates are mesic to dry. The habitats themselves, however, are somewhat humid, usually partially shaded (to open). LOPHOZIA (BARBILOPHOZIA) HATCHERI (Evans) Steph. Plants large, ca. 2-3 mm broad, shoots appearing somewhat julaceous; green to golden- or reddish-brown. Leaves J-k-loted, symmetrical, flat to somewhat undulate, averaging 1.12 times as wide as long, the lateral margins straight but not parallel, leaves narrower at the base than at apex; antical leaf margin usually straight or slightly curved; lobes with straight sides, apices obtuse, acute, or mucronate; distal cells of leaf tip or mucro 10-30 x (16,18)24-88 jam (mean 17 x 47 |0m); middle lobes quadrate or slightly broader or longer, the width ranging from 0.8-1.8 times the length (mean 1.2); postical c i l i a present on leaves, usually 2-5, relatively short, distal cells 8-18 x 20-102(148) |im (mean 12 x 71 |im). Underleaves relatively large, lanceolate-ciliate or bifid-ciliate. Gemmae common and usually abundant, reddish, 2-celled, angular. Sporophytes not observed. Distribution: Gircumboreal, Arctic-Alpine, Bipolar; i n the study area, Dry Northern-Interior. Ecology: Essentially a l l collections were from rock substrates, where the species grows on i t s own humus, needle-litter, or s o i l over boulders and outcrops. Exceptions were l i t t e r at the base of a c l i f f , and humus over s o i l on a streambank. The habitat i s usually a boulder-slope, and the site i s often exposed, occasionally partially shaded i n forest; dry to mesic. LOPHOZIA (BARBILOPHOZIA) LYGOPODIOIDES (Wallr.) Gogn. Plants robust, ca. 4 mm broad, translucent green to golden-green. Leaves (3~)4-lobed, asymmetrical, undulate-crispate, the width 1.2-1.5 (mean 1.3) times the length, narrower at the base than at the apex, lateral margins not parallel, antical margin usually strongly arched; lobes convex-sided, apices mostly mucronate (occasionally acute), d i s t a l c e l l s of leaf t i p or mucro (8)10-20(24) x 40-120 \m (mean 16 x 76 (im), middle lobes much broader than long, the width at the lobe base I.O-3.O (mean 2.0) times the length; postical c i l i a numerous, frequently 5-8, long, d i s t a l c e l l s 8-16 x (32)46-110 [im (mean 12 x 82 |im). Underleaves relatively large, lanceolate-c i l i a t e or b i f i d - c i l i a t e . Gemmae rare, angular, green to reddish, 2-celled (according to the literature; not observed on plants from the study area). Sporophytes not observed. Distribution: Gircumboreal, Arctic-Alpine (?); i n the study area, Dry Northern-Interior. Ecology: Occurring i n forests or on boulder-slopes, primarily on rock substrates, as boulders, where i t i s not a pioneer, but grows over humus, mosses, and l i t t e r . Also on s o i l with humus on streambanks, rarely on decaying logs or l i t t e r over tree (spruce) bases. Substrate mesic to 108 drying, or occasionally with seepage or mist; partially shaded to open. LOPHOZIA Subg. LEI0G0LEA K. Mull. Plants opaque yellowish-green to brownish; growing on calcareous substrates. Medulla of stem entirely homogeneous, lacking a ventral mycorrhizal band. Rhizoids dense, scattered. Leaves bilobed; leaf insertion very succubous to horizontal. Postical leaf base lacking c i l i a . Median leaf c e l l s rather large. Trigones usually large to bulging. Cuticle strongly papillose-striate. Oil-bodies relatively large, pale golden to brownish. Underleaves present i n species i n this f l o r a . Dioicous or paroicous. Perianths long, tubular, essentially nonplicate, contracted at apex to form a beak. Schuster (1969a) discusses the d i f f i c u l t i e s involved i n working with the subgenus Leiocolea. In southwestern B r i t i s h Columbia at least two species are present, and are separated by the following key: 1. Median leaf c e l l s ca. 35~^0 x 45~50 |im; plants lacking gemmae; paroicous L. GILLMANII 1. Median leaf cells ca. 25 x 25~35 P-m; plants usually with 2-celled, golden-brown, smooth, ovoid gemmae, produced on erect, attenuate shoots; dioicous L. HETER0C0LP0S LOPHOZIA (LEIOCOLEA) GILLMANII (Aust.) Schust. Plants light yellow-green to dark green or blackish, 1.5-2.5 mm wide, creeping. Leaf lobes blunt to acute, sinus may be somewhat gibbous. Median leaf c e l l s ca. 35-40 x 45-50 p.m. Trigones moderate to strongly bulging. Cuticle strongly papillose-striate. Oil-bodies pale golden to brownish, 2-6(-10) per c e l l , spherical to ovoid, granular or consisting of minute spherules; 4.0-7.5 M-m i n diameter, or 5-10 x 6-14 |im. Underleaves often obscured by rhizoids, subulate to lanceolate, often with basal c i l i a . 109 Gemmae lacking. Paroicous. Sporophytes found i n August (at low and subalpine elevations). Distribution: Circumboreal - Missing i n Eastern Asia;"in the study area, Widespread (elevation-independent, basic substrates). Ecology: Occurring chiefly on sandy s o i l over calcareous rock outcrops and c l i f f s , including limestone and serpentine; also growing among other bryophytes on l i t t e r and humus on the ground or over decaying logs i n run-off channels or depressions i n the subalpine forest floor. The substrate i s usually mesic to moist with seepage (or spray), and par t i a l l y shaded to open. LOPHOZIA (LEIpGpLEA) HETER0G0LP0S (Thed.) Howe Plants green to golden-brown or reddish, 2.0-2.5 mm wide; shoot apices tending to be erect, especially when gemmiparous. Leaf lobes (obtuse to) acute. Median leaf c e l l s ca. 25 x 25-35 (im. Trigones usually large; c e l l walls usually pigmented. Cuticle strongly papillose-striate. Oil-bodies pale golden to brownish, 2-6 per c e l l , spherical to ovoid, granular, 4.0-7.5 urn i n diameter, or 4.0-7.5 x 5 - H t-im. Underleaves present, ovate-lanceolate, often with c i l i a . Gemmae usually present, at apices of erect, attenuate shoots; 2-celled, red-gold to golden-brown, (14)18-20 x 20-28 jam. Dioicous. Sporophytes not observed. Distribution: Gircumboreal; i n the study area, Dry Northern-Interior (basic substrates). Ecology: Occurring on calcareous, inorganic substrates (e.g., on limestone and volcanic rock), especially on boulders and outcrops (with sand, s i l t , or i t s own humus, occasionally mixed among mosses). Also, 110 occasionally found on (sandy) s o i l and humus on the ground or over roots or decaying logs. The species i s found i n the forest or i n subalpine parkland/meadows, and often near streambanks or waterfalls. Substrate usually mesic to moist (seepage), and partially shaded to open. LOPHOZIA Subg. LOPHOZIA Dum. emend. K. Mull. Plants variable i n size and color. Stem tissues differentiated, i n transverse section, the ventral portion of the medulla becoming mycorrhizal and brownish. Rhizoids scattered. Leaves succubously inserted, 2-lobed. Underleaves absent. Gemmae usually present, 2-celled, angulate. Dioicous or paroicous. 1. Gemmae regularly absent; leaves ± quadrate, sinus broad, shallow; plants usually hygrophytic, t e r r e s t r i a l i n seepy, subalpine meadows L. WENZELII 1. Gemmae usually present, often abundant; leaves variously shaped; plants not hygrophytic, habitats various 2 2. Gemmae gold, reddish-yellow, golden-red, reddish-brown, red, or purplish 3 2. Gemmae colorless, i.e., green . 5 3 . Leaves narrow, longer than wide, the lobes straight, elongated and hornlike, apices often obtuse; plants erect, pure green; gemmae yellow-red to bright brick-red; dioicous . . . . L. LONGIDENS 3 . Leaves at least as broad as long, the lobes not elongated and hornlike, apices usually acute; plants not growing erect, rather ± prostrate-ascending; color, gemmae, and sexual condition various . . . 4 I l l 4. Plants golden-green to gold to golden-brownish (any purplish-red pigmentation confined to ventral stem, and occasionally, also on bases of rhizoids an ventral leaf-bases); oil-bodies ca. 2-11 per c e l l ; gemmae 14-22 p.m long, golden, golden-red, or golden-brown; dioicous L. ALPESTRIS 4. Plants pure green, often with bright purplish-red secondary pigmentation i n the c e l l walls and along the margins of leaves and bracts; oil-bodies numerous, to 20-30 or more per c e l l ; gemmae very large, to 30-40 (im long, bright purplish-red; paroicous L. EXGISA 5. Leaves quadrate, sinus very broad and shallow, descending only to 0.15 x the leaf length; plants t e r r e s t r i a l i n seepy subalpine meadows . . . . . L. WENZELII 5. Leaves usually longer than wide, sinus deeper, 0.2-0.4 x the leaf length; plants not te r r e s t r i a l i n subalpine meadows . . . . . 6 6. Trigones dis t i n c t l y and strongly bulging; oil-bodies ca. 4-l4 per c e l l ; perianth mouth with c i l i a usually 3 -5 ce l l s long; plants v i r t u a l l y restricted to decaying wood . . . . . L. GUTTULATA 6. Trigones usually concave; oil-bodies ca. (ideally) 10-13 per c e l l ; perianth mouth with short teeth, 1-2 cel l s long, often not free for entire length; habitat variable, but plants often growing on rock 7 [L. VENTRIGOSA s. l a t . ] 7. Oil-bodies biconcentric, consisting of minute spherules with one large, central "eyespot," or enlarged, homogeneous spherule L. VENTRIGOSA var. SILVIGOLA 7. Oil-bodies lacking eyespots, appearing granular or consisting uniformly of minute spherules L. VENTRIGOSA var. VENTRICOSA LOPHOZIA (LOPHOZIA) ALPESTRIS (Schleich.) Evans Plants small to medium-sized, 1-2 mm wide; green to golden- or red-brown, with purplish-red coloration occasional on ventral leaf bases. Stems becoming slightly to darker golden- to red-brown, or ventrally purplish-red. Rhizoids abundant, colorless or occasionally purplish-red at the bases. Leaves ovate-quadrate, the width (0.8)0.9-1.0 x the length; bilobed, the lobes triangular, apices obtuse to acute; sinus U-shaped or obtuse, descending to 0.1-0.3 x the leaf length. Median c e l l s ca. 112 15-22(26) x (l4-)20-30 |im; marginal c e l l s quadrate and 14-24 \m long, or slightly rectangular, ca. 12 x 16 |_im. C e l l walls often becoming pigmented, golden to red-gold; trigones lacking or small and concave to slightly bulging; cuticle striate. Oil-bodies 2 - l l ( - l 6 ) per c e l l , usually granular, (or with minute spherules), colorless, usually spherical and 2.5-8.0 \xm i n diameter, or ovoid and 2.5-7-5 x 4-10 [im. Gemmae angular (stellate), 2-celled, red-gold to red-brown or red, ca. 15-25 x 14-26(30) p.m. Dioicous. Sporophytes produced from August through October. Grolle (1976) uses L. sudetica (Nees in Hub.) Grolle for this species. Schuster and Damsholt (1974) support retention of L. alpestris. Distribution: Gircumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. Ecology: Usually growing with s o i l , mosses, humus, or l i t t e r over rock outcrops or boulders. Infrequently, on s o i l and humus on the ground. Substrate usually moist to wet, frequently with.seepage, and open. LOPHOZIA (LOPHOZIA) EXGISA (Dicks.) Dum. Plants variable i n size, smaller forms ca. 350-650 |im wide, other forms larger, to 1760-2000 |_im wide; deep, pure green, becoming tinged with brick-red or wine-red on the upper parts of the leaves and in the c e l l walls. Stem red (-brown) ventrally. Leaves quadrate, the width 0.95-1-10 x the length, 2-lobed, the apices acute; sinus descending to 0.25 times the leaf length. Median cells 30-40 x 40-44 |im. Trigones lacking or small to medium-sized. Cuticle smooth. Oil-bodies 6-20(-30 or more) per c e l l , granular or consisting of minute spherules; spherical and 2.5~7-5 P-m i n diameter, or ovoid-ellipsoid and 2-5-6.0 x 5-10 (im. Gemmae angular, 2-celled, bright brick-red to deep purplish- or wine-red, very large, to 30-40(46) |im long. Paroicous. Perianth mouth purple, with elongate teeth that are free at the ends. Sporophytes found in July, October to November. 113 Distribution: Gircumboreal, Bipolar; in the study area, Relatively Widespread but not oceanic (elevation-independent). Ecology: Growing with mosses, humus, and soil over faces of boulders and outcrop-slopes. Substrate mesic, open to partially shaded. References: Evans 1906a, Schuster I968. LOPHOZIA (LOPHOZIA) GUTTULATA (Lindb. et H. Arn.) Evans = Lophozia (Lophozia) porphyroleuca (Nees) Schiffn. Plants small, 1.0-1.5 mm wide; yellowish-green, often with the postical leaf bases and even the entire leaves becoming red- to wine-colored. Ventral stem red or brownish-red to red-purple. Leaves oblong, the width 0 . 6 0 - 0 . 8 5 x the length; bilobed, the lobes long-triangular, apices obtuse to acute; sinus obtuse to U-shaped, descending to 0 . 2 - 0 . 4 x the length of the leaf. Median cells 20-24(30) x 30-40(54) p.m. Trigones moderately to usually greatly bulging. Cuticle smooth. Oil-bodies 4 - l 4 ( l 8 ) per cell, consisting of minute spherules; spherical, (2.5)3-5 -7.5(9.0) pirn, or ovoid-ellipsoid, 2 . 5 - 6 . 0 ( 8 ) x (3-5)5-15 P-m. Gemmae angular, 2-celled, green, 20 x 20-26 (im. Dioicous. Perianths usually present; perianth mouth with narrow c i l i a formed of 1-3 elongate cells. Sporophytes frequent, May through October (at both low and subalpine elevations). Elaters 8 [in i n diameter. Spores 7-9 pm in diameter. Distribution: Circumboreal; in the study area, Widespread (elevation-independent). Ecology: Primarily a forest species; occasionally found in sites adjacent to the forest, e.g., fields, boulder-slopes, or subalpine parkland. The species grows on decaying wood (logs, stumps, rotting wood), occasionally with humus, mosses, and l i t t e r . On boulder-slopes i t 114 generally grows on rotting wood among boulders, or, rarely, on humus and l i t t e r over boulders; never over bare rock or inorganic substrates. References: Evans 1906a. LOPHOZIA (LOPHOZIA) LONGIDENS (Lindb.) Mac. Plants small, 1.5-2.0 mm wide; dark, pure to yellowish-green; growing ± erect. Leaves narrow, the width O.7O-O.85 " the length, parallel-sided, squarrose; bilobed, the lobes hornlike, apices obtuse to acute; sinus deeply lunate to obtuse. Median leaf c e l l s ca. 25 x 30-35 UJIW Trigones small to slightly bulging. Cuticle smooth. Gemmae angulate, 2-celled, reddish-yellow to bright brick-red. Dioicous. A l l plants i n the study area were s t e r i l e . Distribution: Circumboreal, Arctic-Alpine; i n the study area, Dry Northern-Interior. Ecology: Most frequently occurring among mosses and over humus and l i t t e r on rock (boulders and outcrops). Also growing on completely organic substrates, i.e., on wood (decaying logs and stumps, bases of l i v i n g Douglas f i r and mountain hemlock trees). Substrate dry to mesic, i n open or p a r t i a l l y shaded sites with high humidity. References: Evans 1907b. LOPHOZIA (LOPHOZIA) VENTRICOSA var. SILVICOLA (Buch) Jones Plants as i n L. ventricosa var. ventricosa (see below), but possibly more robust (?)t Primarily differentiated from var. ventricosa by the eyespots on the oil-bodies, i.e., the presence of one much larger homogeneous spherule upon an oil-body which i s otherwise granular or consisting of minute spherules. Oil-bodies i n this variety numerous, (7)10-23 per c e l l , spherical or e l l i p t i c a l , 3'0-7«5 M-m i n diameter or 115 2.5-6.0 x 4-9 urn. Sporophytes found i n April at Deek's Greek. Distribution: Disjunct North America — Western Eurasia; i n the study area, Relatively Widespread but not oceanic (elevation-independent). Ecology: At low elevations, growing on humus, dead wood, or s o i l over rock, on boulders on boulder-slopes, or outcrop-slopes. Once at low elevation on a decaying log i n a marsh (Sunshine Coast). In the subalpine forest, growing on wood, on bases of dead or l i v i n g trees (then mountain hemlock), or on decaying logs. References: Banwell 1949. LOPHOZIA (LOPHOZIA) VENTRICOSA var. VENTRICOSA (Dicks.) Dum. Plants medium-sized, 1-2 mm wide; green or tinged with golden-red, c e l l walls becoming golden-red. Ventral stem red to purplish. Rhizoids abundant. Leaves ovate-quadrate, the width 0.8-1.1 x the length; bilobed, the lobes triangular, apices obtuse-acute; sinus broadly U-shaped, or open and obtuse, descending to 0.20-0.35 x "the length of the leaf. Median ce l l s 18-22(30) x 24-36 (im. Trigones small to bulging, occasionally sinuate. Cuticle smooth. Oil-bodies (2)4-12(-20) per c e l l , (granular to) consisting of minute spherules of uniform size; spherical, (2.5)4-6(7.5) M-m i n diameter, or ovoid-ellipsoid and 2.5-6.0(7-5) x 4-10 |im. Gemmae angular, 2-celled, green, 14-22 x 20-34 (im. Dioicous. Perianth mouth with short teeth, 1-2 c e l l s long, the sides of which are often not free. Sporophytes occasional, May through October at low and subalpine elevations. Distribution: Circumboreal - Missing i n Eastern Asia; i n the study area, Relatively Widespread but not oceanic (elevation-independent). 116 Ecology: Primarily occurring i n rocky places, especially on boulder-slopes, also on outcrops of stream canyons. In such habitats the species grows on s o i l , mosses, i t s own humus, l i t t e r , or rotting wood over the rock substrate. Also occurring i n forests and subalpine parkland, where i t grows on decaying wood (stumps, logs) or, less frequently, on s o i l and humus on the ground or on tree bases (mountain hemlock). Substrate dry to mesic, open (to partially shaded). LOPHOZIA (LOPHOZIA) WENZELII (Nees) Steph. Plants medium-sized to large, 1.5-3'0 J™ wide; light yellow- to golden-green; the shoots having a distinctive growth habit with the apices rolled forward when dry. Stems distinctive, dorsally whitish and opaque, ventrally red or red-gold to red-brown. Leaves quadrate, the width 0.8-1.1 x the length; bilobed, the lobes broadly triangular, apices blunt to somewhat acute; sinus broad, crescent-shaped, very shallow, descending only to 0.05-0.15 x the leaf length. C e l l walls becoming golden; trigones moderate-sized, concave to bulging. Oil-bodies 4-22 per c e l l , granular or with minute spherules; spherical, 2.5-6.5(7-5) m^ i n diameter, or ovoid-ellipsoid, 2.5-6.0 x (5)7«5-10'0 M-iu. Gemmae angular, 2-celled, green; roughly quadrate and 20 ujn on a side, or 16-20 x 22-34 |im, rare. Dioicous. Sporophytes rare, found i n August. Distribution: Gircumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. Ecology: Primarily occurring i n subalpine meadows; occasionally i n subalpine forest or parkland. Generally t e r r e s t r i a l , hygrophytic, growing on the ground over l i t t e r , humus, or s o i l ; occasionally oh l i t t e r , humus, and sand over rocks, boulders, or outcrops. LOPHOZIA Subg. MASSULA K. Mull, emend. Schust. 117 Plants medium to large. Stems, in transverse section appearing flattened, the tissues undifferentiated. Rhizoids scattered. Leaves succubously inserted, 2-3-4-lobed. Leaf cells usually thin-walled, lacking trigones. Oil-bodies generally minute, homogeneous, and numerous, 25-40 or more per c e l l . Gemmae often present. Dioicous. 1. Cells of leaf middle relatively large, to 60 (im or more i n length; gemmae smooth and spherical or ellipsoid; plants occasionally with golden to brick-red secondary pigmentation . . . . LOPHOZIA (new species) 1. Cells of leaf middle small, to 40 \im long; gemmae angulate or lacking; secondary pigments lacking (except i n L. obtusa) . . . . . 2 2. Leaves bilobed, the lobe apices obtuse; plants ± flattened . . L. OBTUSA 2. Leaves 2-3-4-5-lobed, the lobe apices acute; plants rather r u f f l y 3 3. Plants opaque bluish-green; leaves mostly as long as broad; margins of leaf lobes spinose-dentate; perianth mouth with long teeth to c i l i a t e ; widespread at low elevations, common on decaying wood (occasionally extending up into subalpine forest) L. INCISA 3. Plants opaque, whitish- to yellowish-green; leaves broader than long; margins of leaf lobes entire; perianth mouth denticulate, the scattered, short teeth free for only part of their length; subalpine, t e r r e s t r i a l i n seepy meadows, only occasionally descending into subalpine forest (on soil) . . . . L. 0PACIF0LIA LOPHOZIA (MASSULA) INCISA (Schrad.) Dum. Plants medium to large, 1.5-2.5 mm wide; opaque bluish-green, soft, lax, r u f f l y . Leaves as broad as long, polymorphic, mostly 2-4-lobed; lobe margins often wavy, apices acute to acuminate and with teeth. Trigones lacking or small. Cuticle smooth. Oil-bodies numerous, (10-15~)20-40 or 118 more per c e l l ; usually homogeneous, occasionally faintly papillose with rather distinct spherules; spherical to ellipsoid; minute, ca. 2.5-5.0 UJH i n diameter; leaving a blackish residue i n ce l l s of dried plants. Gemmae 2-celled, angulate, green. Dioicous. Perianth mouth c i l i a t e or long-dentate. Sporophytes produced from Ap r i l through August. Distribution: Gircumboreal; i n the study area, Widespread (elevation-independent). Ecology: Usually occurring i n forests, occasionally at the edges of forests, i n ditches or on road banks. Approximately 2/3 to j/k or the collections were from decaying wood, especially rotting logs and stumps. The species also grows on l i t t e r , humus, and sand over rock; on humus and l i t t e r on the ground; on bases of l i v i n g trees (Sitka spruce); and on sandy s o i l banks. Substrates are mostly mesic to moist, par t i a l l y shaded to open. References: Steere 1975-LOPHOZIA (MASSULA) OBTUSA (Lindb.) Evans Plants large to robust, 2-4 mm wide; translucent, light yellow-green to golden to orange-gold, becoming bright brick-red on postical leaf bases; growing i n loose mats. Stems bright brick-red ventrally. Leaves distant to imbricate, very obliquely to horizontally inserted; ± symmetrically bilobed (rarely trilobed), the postical lobe a l i t t l e larger; lobes broad, apices broadly rounded to obtuse; sinus descending ca. 0.30-0.45 x the leaf length. Underleaves present, minute and with c i l i a , or becoming large, lanceolate-ciliate; often obscurred by rhizoids. Median cel l s 20-30 x 26-34 ujn; marginal cells 14-24 x 20-26 \m. Cells thin-walled with trigones small to somewhat bulging; cuticle heavily striate. Oil-bodies ca. 20-30 or more per c e l l , homogeneous to finely granular or 119 "wrinkled," roughly spherical, 1.25-3.75 (im i n diameter. Gemmae 1-2-celled, angulate, green; occasionally formed on mature leaves, whose lobe margins then become eroded. Dioicous. Juvenile archegonia apical-terminal; no perianths or male plants observed. Sporophytes not found. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia; i n the study area, Dry Northern-Interior. Ecology: Growing on needle-litter, mosses, s o i l , or humus on the ground or over rock i n dry coniferous forest or on exposed rock outcrops. Substrate moist to mesic to dry, open (or part i a l l y shaded). References: Muller 1942a. LOPHOZIA (MASSULA) OPAGIFOLIA Culm. Plants medium to large, 1.5-3«0 mm wide; opaque whitish- to yellowish-green. Leaves broader than long, polymorphic, mostly 3-^-lobed; lobes broadly triangular, entire, the apices obtuse or acute. Cell walls thin. Trigones absent or small, rarely bulging. Cuticle smooth. Oil-bodies numerous, 20-35 or more per c e l l ; ± homogeneous, minute, similar to those of L. incisa. Gemmae 2-celled, angulate, green, produced among juvenile leaves at the stem apex. Dioicous. Perianth mouth with scattered, short teeth 1 c e l l long, whose la t e r a l margins are only shortly free from adjacent c e l l s ; occasionally becoming almost entire. Sporophytes produced from August through October. Distribution: Disjunct North America — Western Eurasia, (alpine); i n the study area, S t r i c t l y Subalpine-Alpine. 120 Ecology: Generally occurring in open subalpine meadows, where i t is terrestrial, hygrophytic, growing on moist, seepy humus or soil which is often sandy. Occasionally found in other subalpine sites, on sandy humus and l i t t e r over rock outcrops, or on sandy soil banks along trails, then descending into subalpine forest. References: Schuster 196lb, Lammes 1974. LOPHOZIA (MASSULA) (new species) Plants large, 2-3 mm wide, light golden-green, becoming tinged with reddish pigments. Stem in transverse section ± flattened, fleshy, the cortical cells darker, hyaline medullary cells undifferentiated. Leaves succubous, broader than long, ca. 1520-2000 u,m wide x 1280-1600 |im long, 2~3(-4)-lobed, the lobes broadly triangular, margins entire, lobe apices obtuse to acute; sinus obtuse, margins reflexed, gibbose. Underleaves absent or minute, or occasionally large ({), then to 180-280 x 280-400 \im, spatulate with entire margins and truncate apex. Median leaf cells 24-36 x 34-62 |im. Cell walls thin, becoming orange-gold to reddish-purple. Trigones minute to small but somewhat bulging. Cuticle smooth. Oil-bodies numerous, (l0-)20-30 or more per cell; faintly granular or with individual minute spherules visible, occasionally with eyespots; roughly spherical, minute, 4-5(8) |im in diameter. Gemmae 2(-3)-celled, smooth, spherical to ovoid, plump, green to light golden, 20-26 x 30-44 UJH. Dioicous. Perianths terminal on main shoots, or appearing intercalary with continued growth of the shoot; the mouth denticulate from projecting ends of cells. Sporophytes found in May at low elevations. Elaters red-brown, bispiral, 8-10 \xm in diameter. Spores red-brown, papillose, 16-24 |_im in diameter. This species will be described and illustrated in a future manuscript. Distribution: Endemic to Western North America; i n the study area, Subalpine-Descending. Ecology: Occurring i n subalpine meadows, hygrophytic, growing on sand and gravel (over rocks) on streambanks where the substrate i s wet with flowing water or spray. Also, descending down the mountains along cold rivers and streams. This i s a rather elusive species. LOPHOZIA Subg. ORTHOCAULIS (Buch) Schust. Plants with scattered rhizoids. Leaves 2-, 3-» or 4-lobed, often with postical c i l i a which consist of quadrate c e l l s . C i l i a t e underleaves often present. Cuticle of leaf c e l l s verrucose. Dioicous. Perianths (infrequent) plicate. References: Grolle i 9 6 0 . 1. Leaves predominantly 2-lobed . . 2 1. Leaves 3 -4-lobed 3 2. Plants medium-sized, to 2 mm wide; ( a l l shoots with virtu a l l y a l l leaves 2-lobed); leaves without a tooth on antical base L. KUNZEANA 2. Plants quite small, to ca. 1 mm wide, fuscous-scorched to inky and blackish; (some shoots i n mat with 3~4-lobed leaves); some leaves with a tooth (or an indication of one) on the antical base L. QUADRILOBA (weak shoots) 3. Leaves predominantly J-lohed, postical bases with abundant c i l i a ; antical leaf base entire; plants medium-sized, 1-2 mm wide, golden-green to golden-brown L. FLOERKEI 3. Leaves 4-lobed on well-developed shoots, 2-3(-4)-lobed on weak shoots; postical leaf bases with small c i l i a ; antical leaf base with a tooth or hook on most leaves; plants quite small, to ca. 1 mm wide, dark reddish-brown and becoming fuscous-scorched to inky, blackish L. QUADRILOBA 122 LOPHOZIA (ORTHOCAULIS) FLOERKEI (Web. et Mohr) Schiffn. Plants medium-sized, 1-2 mm wide; golden-green to golden-brown. Leaves predominantly 3-lobed, the lobes very broadly triangular, apices acute; sinuses descending 0.18-0.33 x the length of the leaf; postical bases abundantly c i l i a t e ; antical bases lacking teeth or c i l i a . Underleaves bilobed with 2 lanceolate lobes, c i l i a t e . Median leaf cells 16-24 x (16)24-34 |im; marginal cells 16-24 \m long. Trigones distinct to bulging. Cuticle papillose. Gemmae lacking. Sporophytes not observed. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia; i n the study area, Subalpine-Descending. Ecology: Lophozia floerkei i s the most common species of subg. Orthocaulis i n southwestern Br i t i s h Columbia. It i s essentially subalpine, occurring i n subalpine meadows, parklands, and forests, where i t grows on inorganic substrates (on s o i l and rock outcrops, with or without humus, l i t t e r , or mosses) or organic substrates (with humus over rotten stumps, on rotting wood or logs, on the ground). Substrate mesic to moist (becoming dry when on rock), partially shaded to open. Where i t occurs, the species i s abundant to very abundant. LOPHOZIA (ORTHOCAULIS) KUNZEANA (Hiib.) Evans Plants small to medium-sized, 0.8-2.0 mm wide; green to golden-brown. Leaves concave; predominantly bilobed, the lobes broadly triangular, apices obtuse to rounded, incurved; sinuses descending 0.30-0.44 x the leaf length; postical bases with c i l i a minute or lacking; antical bases without teeth. Underleaves bilobed with 2 lanceolate-ciliate lobes, or variable, often with 1 lanceolate lobe. Median c e l l s 12-18 or 20-24 |im wide x 123 18-34 |im long; marginal c e l l s 14-20 am long. Trigones distinct to minute or lacking. Cuticle verrucose. Gemmae infrequent, 2-celled, angular, golden-green to brownish-red, 14-16 x 16-32 urn. Sporophytes not observed. Distribution: Gircumboreal - Missing i n Eastern Asia, Arctic-Alpine; i n the study area, Dry Northern-Interior. Ecology: At low elevations and at Joffre Creek, growing on s o i l , l i t t e r , and i t s own humus over boulders on boulder-slopes; substrate dry to mesic, pa r t i a l l y shaded to open. At Washington Pass, east of Ross Lake, t e r r e s t r i a l i n moist to wet, open subalpine meadows. References: Evans 1905-LOPHOZIA (ORTHOCAULIS) QUADRILOBA (Lindb.) Evans Plants small, ca. 800-1040 am wide; appearing fuscous or scorched, brownish- to blackish-green and inky. Leaves on weak shoots 2-3-lobed, on robust shoots, 3 -4-lobed; the lobes triangular, apices obtuse to acute; sinuses descending 0.33-0-35 x the length of the leaf; postical leaf base with small c i l i a ; antical base with a tooth or hook on most leaves of well developed shoots. Underleaves mostly lanceolate-ciliate. Median cells 12-24 x 18-34 am; marginal cells 14-20 am long. Trigones small or distinct. Cuticle very strongly verrucose. Gemmae lacking. Sporophytes not observed. This species varies greatly i n form, in the area of this flora as elsewhere i n i t s range. Distribution: Circumboreal - Missing i n Eastern Asia, Arctic-Alpine; in the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on sand over rocks i n a stream below the glacier 124 on Mt. Matier (Joffre Greek T r a i l ) , and i n alpine meadows on Whistler Mountain. Substrate wet, open. LOPHOZIA Subg. PROTOLOPHOZIA Schust. LOPHOZIA (PROTOLOPHOZIA) ELONGATA Steph. Plants light yellow-green, small, 1.0-1.5 mm wide, the leafy portion of shoots 5~9 mm high, the lower portion of the stem becoming leafless, rhizomatous, and tangled i n the bryophyte mat. Stems pale i n color except the ventral cortical cells becoming brownish. In transverse section, stems 9-10 c e l l s high (160-190 (im), 11-14 cells broad (200-230 p.m); a l l c e l l s thin-walled, with no differentiation, variable i n size and shape; cortical cells 14-20 x 18-26 |im i n diameter; medullary cells, 14-24 x 18-30 |im. In surface view, dorsal cortical c e l l s narrow, 60-70 \m long. Rhizoids dense, scattered. Leaves ± quadrate, somewhat distant to approximate on s t e r i l e portions of shoots, succubously inserted; 2(-3)-lobed, the lobes triangular, lobe apices acute; the sinus angulate, rounded, quite deep, descending to 0.35-0.45 x the length of the leaf; leaf margins entire except frequently with a tooth at the antical base. Underleaves distinct under compound microscope, ca. 85-120 p,m long, 30 (im wide at base; lanceolate with a late r a l tooth, or this tooth occasionally becomes a small second lobe. Marginal cells of leaf 14-20 x 18-34 |im. Median leaf c e l l s (14)18-24 x 24-34(48,50) |im. Leaf cells thin-walled, trigones small, concave-sided, distinct. Cuticle smooth to faintly striate. Oil-bodies relatively small and numerous, Massula-like, 12-28 per c e l l , very fa i n t l y papillose, roughly spherical and 2.5-4.0(5.0) [im i n diameter, or occasionally ovoid-ellipsoidal and J-k x 5-6 um. Gemmae lacking. Paroicous. F e r t i l e shoots with 2 to 4 or 5 sets of bracts below perianth. Male bracts similar to leaves but larger, somewhat more 125 elaborate, with an antical tooth. Antheridia occurring l(-2) per bractj the body ca. 78-90 x 104-110 p.m; stalk uniseriate, 10-14 \m i n diameter., Female bracts larger and more elaborate than leaves, 3-^-l°bed, occasionally with accessory teeth or lobes on lateral margins, the lobes somewhat undulate. Bracteole 1-2-lobed, occasionally with accessory teeth on l a t e r a l margins. Perianth terminal on main shoots, to 2.5 mm or more in length, obovate, plicate d i s t a l l y . Perianth mouth divided into several lobes, the lobes irregularly dentate-ciliate; the teeth 10-14 x 36-48 |im, 1- or 2-celled. Immature sporophytes present; seta 8 ce l l s i n diameter, with 18 epidermal and ca. 21 inner rows of ce l l s . Lophozia elongata i s a very rare plant. The specimen from the Brooks Peninsula appears to possess the features which distinguish the species elsewhere. Results of further study of this specimen w i l l be discussed i n a future manuscript. Distribution: Western North America — Greenland — Western Europe (rare, sporadically distributed northern species; previously known from Norway, Sweden, Yugoslavia (?), West Greenland); i n the study area, Rare or Restricted (plants known only from the Brooks Peninsula, northwestern Vancouver Island). Ecology: Growing on sand over c l i f f face, wall of ravine on ocean beach. Substrate mesic, i n pa r t i a l shade. References: Schuster and Damsholt 1974. TRITOMARIA Schiffn. Plants with 3-lobed leaves, succubously inserted; lacking c i l i a on postical leaf bases. Underleaves lacking. References: Schuster 1958c. 126 1. Leaves symmetrically 3-lobed 2 1. Leaves very asymmetrically 3-lobed 3 2. Plants soft, lax i n appearance; leaf lobes blunt to rounded; leaf insertion dorsally arched toward stem apex; ce l l s of leaf bases strongly elongated, 4-8 x as long as wide T. POLITA 2. Plants s t i f f i n appearance; leaf lobes sharp; leaf insertion dorsally transverse; basal leaf c e l l s not strongly elongated T. SCITULA 3. Leaves wider than long; plants large, 1.5-3«0 mm wide . . T. QUINQUEDENTATA 3. Leaves narrow, longer than wide; plants smaller, 1.0-1.5 mm wide 4 4. Gemmae smooth, ovoid-ellipsoid; median leaf c e l l s smaller, 12-26 u.m long T. EXSECTA 4. Gemmae angular; median leaf c e l l s larger, I8-36 urn long . . . . T. EXSEGTIFORMIS TRITOMARIA (TRITOMARIA) EXSEGTA (Schrad.) Loeske Plants small, 1.0-1.5 mm wide; pale green; erect-ascending. Leaves asymmetrically 3-l°bed, longer than wide. Median leaf c e l l s 12-26(30) |im long. Gemmae red-brown, smooth, ovoid-ellipsoid, 2-celled. Dioicous. Plants from the study area were st e r i l e . Distribution: Gircumboreal, Arctic-Alpine (?); i n the study area, Rare or Restricted (distribution not well understood). Ecology: Growing i n humid sites, on a dry cedar log on a lakeshore, and on trunks of crabapple trees on a streambank near a large lake. Substrate dry to mesic, partially shaded. TRITOMARIA (TRITOMARIA) EXSEGTIFORMIS (Breidl.) Loeske Plants virtually identical to T. exsecta, differing i n the larger median leaf cells, I8-36 |im long, and in the gemmae, which are 2-celled, red-brown, and angular. Dioicous. Only s t e r i l e plants were found. Distribution: Circumboreal - Missing i n Eastern Asia, Arctic-Alpine (the species s. l a t . ) ; i n the study area, Dry Northern-Interior. Ecology: Growing on humus over rotting logs, boulders, and i n outcrop crevices. Substrate dry to mesic, partially shaded. TRITOMARIA (SAGCOBASIS) POLITA (Nees) Joerg. Plants medium to large, 2-3 mm wide, green, appearing soft and lax; said to have a greasy lustre. Dorsal cortical c e l l s of stem elongate, thick-walled, 4-8 x as long as wide. Leaves symmetrically 3-l°t>ed, wider than long. Oil-bodies 2-9 per c e l l , consisting of minute spherules, spherical and 4-6 urn i n diameter. Gemmae not observed; said to be rare. Dioicous. Plants from the study area were s t e r i l e . Distribution: Disjunct North America — Western Eurasia, (arctic-alpine); i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on s o i l over streambanks, boulders, and c l i f f s i n subalpine locations. Substrate mesic to wet with flowing water, open to parti a l l y shaded. TRITOMARIA (TRITOMARIA) QUINQUEDENTATA -(Huds.) Buch Plants medium-sized to large, 1.5-3«0 mm wide, often rather lax, growing i n prostrate tufts, yellowish-green or brownish. Leaves asymmetrically 3-l°t>ed, wider than long. Gemmae rare i n this region, angular, yellowish. Dioicous. Sporophytes rare, produced i n August and September (subalpine s i t e s ) . 128 Distribution: Gircumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. Ecology: Typically occurring i n subalpine meadows, parkland, or forest (along streams), where i t grows on humus and s o i l either on the ground or over rock (boulders, c l i f f s , or outcrops). Occasionally descending to lower elevations i n humid l o c a l i t i e s , along cold streams or near waterfalls. Substrate mesic to moist or wet, open or partially shaded. TRITOMARIA (TRITOMARIA) SGITULA (Tayl.) Joerg. Plants medium-sized, 1-2 mm wide, bright green. Leaves symmetrically 3-lobed, longer than wide; the lobe apices obtusely, bluntly pointed, lobes somewhat hornlike. Gemmae red, angulate, 1 -2-celled. Dioicous. Plants i n the study area were st e r i l e . Distribution: Circumboreal - Missing i n Eastern Asia, Arctic-Alpine (?); i n the study area, Rare or Restricted (distribution not yet well known). Ecology: Growing among other bryophytes on calcareous rock (including limestone and basaltic rock); also found on a decaying log below a wet limestone c l i f f . Substrate mesic to moist, or wet with seepage; open to par t i a l l y shaded. References: Evans 1912. Subfamily JAMESONIELLOIDEAE JAMESONIELLA (Spruce) Schiffn. JAMESONIELLA AUTUMNALIS (DC) Steph. Plants medium-sized to large, ca. 2 . 0 - 2 . 5 mm wide; medium green to golden to bright red, becoming dark purplish-red when dry. Leaves 129 orbicular-ovate. Underleaves absent or vest i g i a l and consisting of minute uniseriate filaments. Trigones absent or small, usually concave (to slightly bulging). Cuticle smooth to punctate. Oil-bodies 6-16 per c e l l , granular, usually ellipsoidal, 4.0-7.5 x 5.0-12.5 |om, occasionally spherical and 4.0-7-5 M-m.. Asexual reproduction lacking. Dioicous ("normally" according to the literature). Bracteole large, divided into narrowly lanceolate segments. Perianth cylindrical, narrowed, plicate i n upper part; perianth mouth long-ciliate. Sporophytes found from late winter to spring. Distribution: Circumboreal; i n the study area, Relatively Widespread but not oceanic ( s t r i c t l y low-elevation). Ecology: Growing primarily on decaying logs or stumps or rotting wood; also on humus and among mosses over rock outcrops or boulders; rarely on trees (crabapple). Substrate mesic to moist, partially shaded to open, i n humid locations, especially i n forests. References: Grolle 1971. Subfamily JUNGERMANNIOIDEAE JUNGERMANNIA L. emend. Dum. Plants prostrate i n growth, with apices of shoots ascending. Stems, i n transverse section, with cortex l i t t l e differentiated from medulla; medulla ± homogeneous. Rhizoids scattered along ventral stem. Branches occasional, including lateral-terminal and lateral-intercalary (normally from lower halves of leaf axils) types. Leaves entire, unlobed (though occasionally refuse at apex), succubously inserted. Underleaves lacking (except on erect, gemmiparous shoots of J. leiantha). Leaf cells with small to bulging trigones; cuticle smooth to striate. 130 Dioicous or paroicous. Female bracts entire, similar to leaves; bracteole absent. Perianth distinct, ± immersed to longly exerted. In this study, the genus Jungermannia i s used i n the broad sense, following Amakawa (1959, 1960a), Vana (1973a, 1973b, 1973c, 1974a, 1974b, 1975a, 1975^), Inoue (1974a, 1976), and Grolle (1976). Schuster (1969a) separates species presented here under Jungermannia into two genera. Jungermannia i s narrowly construed to include only J. lanceolata (= J. leiantha i n this f l o r a ) . The remaining species are placed i n the genus Solenostoma Mitt. The genus Jungermannia has always given r i s e to taxonomic problems. The essential facts of the matter are as follows. Jungermannia was designated by Linnaeus i n 1753 "to include most leafy Hepaticae. Since that time the genus Jungermannia has become generally associated with entire-leaved species of Nardia, Jungermannia (J. lanceolata), and Solenostoma (Schuster 1969a: 835). Schuster (1969a: 835) regards the plant Jungermannia lanceolata as belonging to a separate genus from plants he places i n Solenostoma, largely on the basis of l) i t s unique, smooth perianth, at the apex of which i s a short beak recessed i n a slight depression, and 2) i t s a b i l i t y to produce fasciculate gemmae from the leaf margins. Schuster (1969a: 836) points out that the name "Jungermannia  lanceolata," as applied to this particular plant, "has been i n continuous use,'in exactly our present sense, since 1797. It i s almost unique among older species of Hepaticae i n lacking any synonymy, indicating a singularly even and uneventful history." Unfortunately, the Linnaean J. lanceolata was apparently a composite species, none of whose components were actually the plant long known by that name. The plant " J . lanceolata" did not have a technically correct diagnosis or epithet u n t i l very recently, when Grolle 131 (1966) described i t as Jungermannia leiantha Grolle. Vana (1973c) and Grolle (1976) treat Jungermannia leiantha as belonging to the same genus with plants that Schuster (1969a) places i n Solenostoma. I f J. leiantha were to be segregated as a separate genus under this system of a broad treatment of the genus Jungermannia, i t would have to be placed i n Liochlaena Nees i n Gott., Lindenb. et Nees, of 1845. I f Schuster's system and the genus Solenostoma were followed i n this f l o r a , an additional problem would arise i n the need to create new combinations for Jungermannia confertissima (synonym = Solenostoma  l e v i e r i ) , J. exsertifolia (synonym = Solenostoma cordifolium), and J. hattoriana. Jungermannia hattoriana was previously known only from Japan. Jungermannia confertissima Nees (1833) has recently been shown (Vana 1974b) to be an earlier name for Solenostoma l e v i e r i , the epithet of which f i r s t appeared i n Nardia l e v i e r i Steph. (I892). Jungermannia exsertifolia Steph. was described i n 1927, but must replace the earlier J. cordifolia Hook. (1812) since this was apparently an illegitimate synonym of J. cordifolia Brotero (1804), which apparently referred to a species of Scapania (Vana 1973b). In summary, then, for the purposes of this present flora, i t seemed most practical, at least, to follow use of Jungermannia i n a broad sense. The resulting names for species w i l l seem strange and unfamiliar to North Americans accustomed to using Schuster's works on Hepaticae. Therefore, after each species name given as a Jungermannia, the synonym as used by Schuster (1969a) i s also given. Hopefully, this w i l l allow the user of this f l o r a to recognize familiar species names while simultaneously being made aware of synonyms used on other continents. In keeping with the format decided upon for this flora, genera and subgenera are arranged alphabetically, i n spite of the fact that this 132 sequence leaves Jungermannia leiantha (in subg. Liochlaena) uncomfortably out of place among species more familiarly grouped as Solenostoma. Species of the genus Jungermannia occurring i n southwestern B r i t i s h Columbia are separated by the following keys: KEY I: PLANTS STERILE 1. Leaves ± rectangular; (perianths completely smooth, tubular, abruptly contracted to a short beak which i s recessed i n a shallow depression); plants growing on decaying wood, logs . . J. (LIOCHLAENA) LEIANTHA 1. Leaves orbicular, ovate, or reniform; (perianths various, but never smooth and tubular nor with a recessed beak); plants growing on s o i l or rock, most species hygrophytic 2 2. Rhizoids reddish-purple 3 2. Rhizoids colorless or pale brownish . 4 3. Plants odorless; shoots much branched, with abundant flag e l l a ; oil-bodies appearing lik e grape-clusters; (dioicous); hyperoceanic J. (PLECTOCOLEA) HATTORIANA 3. Plants having a strong turpentine-like odor; shoots lacking fl a g e l l a ; oil-bodies consisting of minute, uniform spherules; (paroicous); plants common i n subalpine meadows or descending to lower elevations along cold streams . . J. (PLECTOCOLEA) OBOVATA 4. Plants with bright red-purple pigments; leaves orbicular, bordered by a row of enlarged, thick-walled cells; generally growing on s o i l i n exposed, disturbed sites, e.g., roadsides J. (SOLENOSTOMA) RUBRA 4. Plants olive- to brownish-green to blackish; leaves not bordered; generally growing along streams 5 5. Leaves as broad as, or broader than long 6 5. Leaves ovate-elongate, longer than broad 7 Plants small, ca. 1.5 mm wide, resembling Nardia scalaris; leaves orbicular to reniform; cuticle smooth;(paroicous); with a Northern-Interior distribution i n the study area . . . . . . . . ' J. (SOLENOSTOMA) CONFERTISSIMA Plants large, 2.0-4.5 mm wide; leaves (orbicular to) broadly triangular, the width just above the leaf base equal to the length of the leaf; cuticle striate; (dioicous); common i n streamlets i n subalpine meadows . . . J. (JUNGERMANNIA) EXSERTIFOLIA Plants growing on limestone or other calcareous rock; (dioicous) J. (JUNGERMANNIA) ATROVIRENS Plants occurring on sandstone or other siliceous rock; (paroicous) J. (JUNGERMANNIA) PUMILA KEY II: PLANTS FERTILE Plants dioicous 2 Plants paroicous 5 Shoots much branched, with abundant postical fl a g e l l a ; rhizoids reddish-purple; oil-bodies appearing like grape-clusters J. (PLEGTOCOLEA) HATTORIANA Shoots lacking fla g e l l a ; rhizoids colorless or pale brownish; oil-bodies granular or consisting of minute spherules 3 Plants bright green to bright reddish; leaves orbicular, bordered with a row of greatly enlarged, thick-walled marginal c e l l s J. (SOLENOSTOMA) RUBRA Plants olive-green to purplish-brown or blackish; leaves not bordered 4 Plants small, 1-2 mm wide; leaves ovate, longer than broad J. (JUNGERMANNIA) ATROVIRENS Plants large, 2.0-4.5 mm wide; leaves usually broadly triangular (to ± orbicular), the width just above the base equal to the length J. (JUNGERMANNIA) EXSERTIFOLIA 5. Leaves ± rectangular; perianths completely smooth, tubular, abruptly contracted to a short beak which is recessed in a shallow depression; plants growing on decaying wood, logs . . . '. J. (LIOCHLAENA) L E I A N T H A 5. Leaves ± orbicular, ovate, or reniform; perianths various, but never smooth and tubular nor with a recessed beak; plants growing on soil or rock, mostly hygrophytic 6 6. Plants with red-purple pigments; rhizoids purple; plants with a strong turpentine-like odor J. (PLECTOCOLEA) OBOVATA 6. Plants olive-green to brownish-black; rhizoids colorless to pale brownish; plants without a distinct turpentine-like odor . . . 7 7. Leaves ± orbicular to reniform; cuticle smooth; (plants resembling Nardia scalaris) J. (SOLENOSTOMA) CONFERTISSIMA 7. Leaves ovate-elongate; cuticle striate . . J. (JUNGERMANNIA) PUMILA JUNGERMANNIA Subg. JUNGERMANNIA = Solenostoma Subg. Luridae (Spruce) K. Mull. Plants olive-green or blackish-green. Stem tissues not specialized. Rhizoids confined to stem. Leaves generally longer than broad. Asexual reproduction lacking. Perianth emergent to 0.6 x i t s length beyond the bracts, fusiform to slenderly obovoid, often slightly compressed, with 1 or 2 deep antical sulci, distally plicate but the plicae not sharp. Cells of perianth similar to median leaf cells. Perigynium lacking. JUNGERMANNIA ATROVIRENS Dum. = Solenostoma atrovirens (Schleich.) K. Mull. Plants small to medium-sized, ca. 1200-2000 [im wide, olive- to brownish-green to blackish. Rhizoids pale brownish. Leaves ovate-elongate. Median leaf cells (14) 16-22 x (18)20-30 \xrn-, marginal cells 14-20(22) pun long. Cell walls yellow-gold to orange. Trigones small to 135 slightly bulging, distinct; cuticle striate, at least at the leaf base. Oil-bodies 2-3(5) VeT c e l l , f a i n t l y granular to homogeneous; spherical and 4.0-7.5 \IM i n diameter, or elongate and 4-6 x (5)9 -13(l5) M-m. Dioicous. Perianth ca. 0.5 emergent, bracts not sheathing; cylindrical, with several plicae at apex; mouth cells with tips more than l/2 free. Sporophytes observed once, i n May. Distribution: Circumboreal, Arctic-Alpine; i n the study area, Widespread (elevation-independent, basic substrates). Ecology: Occurring on calcareous rock, boulders, and c l i f f s , i n forests by rivers and streams. Substrate moist to wet, partially shaded to open. JUNGERMANNIA EXSERTIFOLIA Steph. = Solenostoma cordifolium (Hook.) Steph. Plants large to robust, 2.0-4.5 mm wide, olive-green to (purplish-) brown to blackish, lax. Leaves ovate to heart-shaped or broadly triangular; as broad just above base as they are long. Cell walls orange-gold; trigones lacking; cuticle striate, at least at the leaf base. Median cel l s 16-26 x 26-48 (am, or usually larger, and 20-36 x 30-66 \xm; marginal c e l l s 18-30(-38) u.m long. Oil-bodies mostly 2 per c e l l , varying to 5( -7); consisting of minute spherules or faintly granular to almost homogeneous; spherical, 4-10 |im i n diameter, or ellipsoid-ovoid, 4-9 x 6-15 [im. Dioicous. Perianths long-emergent (to 0,5 x the perianth length), plicate i n upper portion; mouth cells fingerlike, more than l/2 free. Bracts not sheathing. Sporophytes observed on plants collected i n late August (subalpine). Vana (1973^ 0 recognizes two subspecies for this species: subsp. 136 cordifolia, from North America, Europe, and across northern Asia, and subsp. exsertifolia, from Japan. Plants from B r i t i s h Columbia examined for this f l o r a seem to f i t into subsp. cordifolia as defined by Vana. Inoue (1974a: 180), states i n his notes on Jungermannia cordifolia Hook., ". . . i n North America I have observed several times plants which quite agreed with Japanese plants . . . i f there are more prominent differences between the populations i n Europe and Japan, then the North American populations must go into the same category as the Japanese populations." Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (alpine); i n the study area, Subalpine-Descending. Ecology: Growing on rock (occasionally with sand) i n streams and streamlets, almost always i n subalpine meadows or coming down into subalpine forest; infrequently, descending to low elevations. Substrate wet, part i a l l y or to t a l l y submerged i n flowing cold water; open or partially shaded i n forests. JUNGERMANNIA PUMILA With. = Solenostoma pumilum (With.) K. Mull. Plants small to medium-sized, 1-2 mm wide; green to golden- (or ' purplish-) brown. Rhizoids colorless. Leaves elongate-ovate. Median c e l l s of leaves 12-26 x 20-40 |im; marginal ce l l s (16)18-26 |_im long. Cell walls colorless or becoming golden; trigones variable, from minute to small to somewhat bulging; cuticle striate. Oil-bodies 2-5(6) per c e l l , granular or with minute spherules; spherical and 3-0-7.5 M-m i n diameter, or elongate and 4-8 x 5-14 \xm. Paroicous. Perianth long-emergent, plicate at very apex; mouth with finger-like c e l l s , ends free. Bracts not sheathing. Sporophytes not found. 137 Distribution: Gircumboreal; i n the study area, Rare or Restricted (distribution poorly understood, too few collections). Ecology: Occurring on siliceous rock (e.g., sandstone) i n streams, on streambanks, or on canyon walls. Substrate moist to wet, with seepage or flowing water for at least part of the year; open to part i a l l y shaded. JUNGERMANNIA Subg. LIOGHLAENA (Nees) S. Arn. = Jungermannia (Ruppius) L. (Schuster 1969a) Plants green (to brownish). Stem tissue not specialized. Rhizoids confined to stem. Leaves ± rectangular to ovate. Asexual reproduction possible by means of fasciculate gemmae from apices of erect, attenuated shoots; Schuster (1969a) states that this i s rarely present. No gemmae were observed on plants from southwestern B r i t i s h Columbia. Perianth cylindrical, smooth, tubular, not at a l l plicate; the apex suddenly contracted to a short beak, which i s recessed i n a shallow depression. Perigynium lacking. 0 JUNGERMANNIA LEIANTHA Grolle = Jungermannia lanceolata L. emend. Schrad. Plants large, 2.5-4.0 mm wide, light yellow-green to red-gold to red-brown. Leaves ± rectangular, parallel-sided, the apex rounded or slightly emarginate. Underleaves absent. Trigones distinct, bulging. Cuticle d i s t i n c t l y striate. Oil-bodies (4)8-15 per c e l l , granular, ovoid and 5-0-7.5 x 6-14 urn, or spherical and ca. 5 _ 8 u.m in diameter. Asexual reproduction lacking on plants from the study area. Paroicous. Female bracts similar to leaves. Perianth tubular and smooth, d i s t i n c t l y contracted at top into beak which i s recessed i n a depression; c e l l s of perianth mouth (beak) have ends protruding. 138 Sporophytes found on almost a l l specimens, Ap r i l through November. Distribution: Disjunct North America — Western Eurasia; i n the study area, Widespread (elevation-independent). (See Grolle and Vana I969). Ecology: Generally growing on decaying logs, or on rotting wood or roots on the ground i n moist forests; also found on a peaty bank i n a swamp. Substrate mesic to moist, usually pa r t i a l l y shaded. JUNGERMANNIA Subg. PLECTOCOLEA (Mitt.) Amak. = Solenostoma Subg. Plectocolea Mitt. Plants bright or light green with reddish-purple secondary pigments. Stems ( i n transverse section) usually with the cortical 1-2 c e l l layers sharply differentiated from the medullary c e l l s . Rhizoids originating from the ventral stem and occasionally also arising from postical bases of leaves. Leaves rotundate to ovate. Asexual reproduction lacking. Perianth reduced, immersed within closely sheathing bracts, or only emergent to 0.35 x i t s length; ovoid, contracted to a slender, but not beaked, mouth. Cells of perianth narrow, elongated, 2.5-5-0 x as long as wide. Perigynium well developed, shortly tubular. References: Amakawa 1960b. JUNGERMANNIA HATTORIANA (Amak.) Amak. Plants 1.0-2.5 mm wide, but quite long (to 2.5 cm) so appearing rather large; deep green, with leaf bases becoming reddish-purple. Shoots much branched, with abundant, often strong f l a g e l l a . Rhizoids reddish-purple. Leaves orbicular to ovate, 0.9-2.8 mm wide, 1.1-2.8 mm long. Median leaf c e l l s 20-42 x 30-66 (im; marginal cells (l2-)20-26 x (20)22-34 \m; basal c e l l s (25-)32-42(-62) x 50-130 (im. Cell walls becoming pinkish 139 to red. Trigones lacking or small but distinct. Cuticle striate. O i l -bodies 2(-4) per c e l l , consisting of various sized globules or grape-clusters, colorless; spherical (7.5)8.5-14.0 u.m i n diameter, or ellipsoid and 6.0-12.5 x 10-19 |im. Dioicous. Perianth l i t t l e exerted. Sporophytes not found. This hepatic, upon f i r s t microscopic examination, immediately appeared different from any species of Jungermannia previously reported from North America. In southwestern B r i t i s h Columbia, J. obovata also has reddish-purple rhizoids, but i t di f f e r s from the present species i n i t s sexual condition (paroicous, rather than dioicous). The presence of abundant strong f l a g e l l a and grape-cluster oil-bodies (oil-bodies consisting of relatively large, homogeneous spherules of various sizes) places these plants into Amakawa's (1960a) Flagellata Group of Jungermannia subg. Plectocolea. S. Hattori and H. Inoue (pers. comm.) believe that the B r i t i s h Columbia plant i s Jungermannia hattoriana. The sizes for leaves, leaf c e l l s , and oil-bodies i n our plants are larger than those given by Amakawa (1960a: 2l) for Japanese specimens. The plants from British Columbia, at least those from the Brooks Peninsula, also have stronger f l a g e l l a than appear to be present i n J. hattoriana of Japan. Three populations of this species are now known from southwestern British Columbia (see distribution map, Appendix C). T. Amakawa (pers. comm.) believes he has seen similar plants belonging to the Flagellata group from the Prince Rupert area, northward on the coast of Br i t i s h Columbia. This species should be sought i n the future on the Queen Charlotte Islands and northward on the mainland coast, and further comparisons should be made with Japanese plants. Distribution: North Pacific; i n the study area, Hyperoceanic. New 140 to North America; previously known only from Japan. Ecology: Growing on sand over rock in streams. In Japan this species is found "on wet rocks or gravels by streams in the subalpine region, 1700-2400 m.s.m." (Amakawa 1960a: 21). JUNGERMANNIA 0B0VATA Nees = Solenostoma obovatum (Nees) Schust. Plants medium (to large), 1.5-2.5 mm wide; green to golden-brown or becoming red-purple to blackish; with a strong turpentine odor when fresh and wet, or even when dry plants are remoistened. Rhizoids red-purple, occasionally originating from abaxial leaf bases. Leaves orbicular-ovate, the apex often emarginate. Median leaf cells 20-30 x 26-40(56) (am. Cell walls occasionally purplish. Trigones small to slightly bulging; cuticle striate. Oil-bodies 2-5 per cell, consisting of minute spherules; spherical, 3'5_5'0 |im in diameter, or ovoid-ellipsoid, 5 _9 x 10-15 \xm. Paroicous. Perianth immersed, bracts sheathing; plicate in upper part; cells of mouth with only the tips free; cells of lower portions of perianth elongate. Sporophytes found in September (subalpine). Distribution: Disjunct North America — Western Eurasia, (alpine); in the study area, Subalpine-Descending. Ecology: Generally growing on sand with humus or moss over rocks and outcrops, on bare rock, or on soil and gravel; occasionally found on decaying logs. Primarily subalpine, where i t occurs on outcrops and banks or rivulets and pools in meadows; occasionally descending to low elevations along cold streams and near waterfalls. Substrates moist to wet, often in flowing water; open to partially shaded. 141 JUNGERMANNIA Subg. SOLENOSTOMA (Mitt.) Amak. = Solenostoma Subg. Solenostoma Mitt. Plants bright green to brownish or reddish. Stem tissue unspecialized. Rhizoids arising from ventral .side of stem, and frequently also from abaxial faces of leaves and bracts. Leaves orbicular to reniform. Asexual reproduction lacking i n moct species. (Endogenously formed gemmae occur i n J. caespititia Lindenb., which does not occur i n B r i t i s h Columbia.) Perianth 0.7 x i t s length emergent beyond the bracts; deeply 4 -5-plicatej abruptly contracted d i s t a l l y into a short beak. Cells of perianth ± similar to median leaf c e l l s . Perigynium lacking or vest i g i a l . JUNGERMANNIA GONFERTISSIMA Nees = Solenostoma l e v i e r i (Steph.) Steph. Plants small, ca. 1.5 mm wide; green to golden-brown, to purplish; resembling Nardia scalaris i n appearance. Rhizoids colorless to pale brown, occasionally occurring on postical leaf-bases. Leaves orbicular to reniform. Median leaf cells 22-32(36) x 26-40(46) [im; marginal c e l l s 18-30 |im long; basal c e l l s 22-44 x 40-50(-60-80) (im. Cell walls colorless, golden, or becoming pinkish-purple at leaf bases. Trigones small but distinct and somewhat bulging. Cuticle smooth (occasionally faintly striate at base of bracts). Oil-bodies 2-7(8) per c e l l , f a i n t l y granular; spherical and 5 - 8 (im i n diameter, or elongate-ellipsoidal and 5 _8 x 7.5-16.0 (im. Paroicous. Perianth slightly to d i s t i n c t l y short-beaked; plicate d i s t a l l y . Sporophytes found i n June at Birkenhead Lake. Epidermal c e l l s of capsule wall with nodular thickenings, 4-5 on long walls, 0-2 on short walls. Inner ce l l s of capsule wall with numerous sharply defined complete 142 semiannular bands. Elaters 8-10 p,m i n diameter, bispiral, the spirals 2-3 um wide. Spores 16-18 (im i n diameter. Jungermannia confertissima (Solenostoma l e v i e r i ) i s similar to, and often d i f f i c u l t to distinguish from, J. sphaerocar-pa Hook., or Solenostoma  sphaerocarpum (Hook.) Steph. Schuster (1969a: 962) presents a table of characteristics by which the two species can be separated. The plants from southwestern B r i t i s h Columbia have, for the most part, features of J. confertissima, but the spore size (spores from previously unopened capsules) i s small, i n the range given for J. sphaerocarpa. These small spores, however, do f i t into the range of 15-22(-28) \m given by Vana (1974b: 372) for J. confertissima. Also, Vana (1974b: 373) notes an ecological difference for the two species, with J. confertissima growing on basic substrates, and J. sphaerocarpa. on siliceous substrates. On that ecological basis alone, one would expect the plants from southwestern Br i t i s h Columbia to belong to J. confertissima. Distribution: Circumboreal (?), Arctic-Alpine; i n the study area, Dry Northern-Interior. Ecology: Growing on sand or sand over rock, on the banks of mountain streams i n the forest. Substrate moist to wet, partially shaded to open. References: Muller 1942a, Paton 1970, Vana 1974b. JUNGERMANNIA RUBRA Gott. ex Und. = Solenostoma rubrum (Gott.) Schust. Plants small, ca. 1.0-1.5 mm wide, with distant leaves, or larger with leaves approximate; bright green to purplish-red. Rhizoids colorless or yellowish, not originating from abaxial leaf bases. Leaves orbicular, with a border of somewhat enlarged cells with thickened walls. Median 143 leaf c e l l s 20-34(42-50) x 28-40(48-70) |om. Trigones small, distinct to bulging. Cuticle striate. Oil-bodies (1)2-4 per c e l l , granular or consisting of minute spherules? spherical and (2)4-5 |im i n diameter, or elongate, 4-6 x 7-5-12.5 |im. Dioicous. Perianths common, bright purplish-red, emergent to about 0. 3 or 0.4 x the length, 4-5-plicate i n upper l/2; c e l l s of mouth elongate-crenulate with tips free. Sporophytes frequent, from March through May. Distribution: Endemic to Western North America; i n the study area, Widespread (essentially low-elevation). Ecology: Frequent on consolidated sand, l i t t e r , and gravel i n roads and t r a i l s , on banks and i n ditches. Substrate mesic to moist, becoming wet where there i s seepage; usually open. NARDIA S. Gray Plants small to medium-sized. Stems fleshy. Rhizoids scattered on ventral stem. Leaves entire or bilobed, succubously inserted. Underleaves distinct, lanceolate. Asexual reproduction lacking. Gynoecia terminal on main shoots. Female bracts closely sheathing the short, immersed perianth. Perigynium present, fleshy. References: Evans 1937; Amakawa 1959; Inoue 1971a, 1971b; Vana 1973a, 1976a. 1. Leaves clearly bilobed; oil-bodies homogeneous or becoming segmented; (occurring i n subalpine meadows) N. JAPONICA 1. Leaves entire (occasionally emargihate); oil-bodies various . . . . 2 2. Plants large, 2-3 mm wide, appearing strongly compressed lat e r a l l y ; leaves broadly reniform; (subalpine, uncommon) . . . N. COMPRESSA 2. Plants smaller, 1.0-2.5 mm wide, not appearing strongly l a t e r a l l y compressed; leaves ± orbicular . . 3 144 3. Plants green to golden-brown; leaves emarginate or entire; oil-bodies consisting of minute spherules; subalpine . N. GEOSGYPHUS 3. Plants green, often with red pigments; leaves entire, broadly rounded; oil-bodies homogeneous-segmented; occurring primarily at low elevations N. SCALAR!S NARDIA COMPRESSA (Hook.) S. Gray Plants large to robust, 2-3 mm wide, becoming bright red. Leaves broadly reniform, apices entire. Underleaves small, but present along entire stem. Oil-bodies 1-2(3) per c e l l , homogeneous or becoming segmented; spherical, ca. 6 \m i n diameter, or ovoid-ellipsoid, then to 8 (im wide x up to 14 \m long. Dioicous. Plants from the study area were st e r i l e . Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (alpine); i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on humus or s o i l on banks or slopes i n subalpine meadows. Substrate moist to wet with seepage; open. NARDIA GEOSGYPHUS (De Not.) Lindb. Plants small to medium, ca. 1.0-2.5 mm wide; green to warm golden-brown. Leaves orbicular to shortly reniform, the apices entire to emarginate. Underleaves distinct along entire stem. Oil-bodies (1)2-4(5,6) per c e l l ; consisting of minute spherules, occasionally appearing lik e grape-clusters; spherical and (5)6-8 |im i n diameter, or ovoid-ellipsoid and 4.0-7.5 x 7.0-17.5 M-m. Paroicous. Sporophytes infrequent, found i n October. Distribution: Disjunct North America — Western Eurasia; i n the study area, S t r i c t l y Subalpine-Alpine. 145 Ecology: Growing on s o i l and humus on the ground or over rock outcrops, on slopes, rivulet banks, and lake shores i n subalpine meadows. Substrate usually moist to wet with seepage; open. References: Evans 1907b. NARDIA JAPONICA Steph. Plants small, 440-960 (im wide, 7-15 mm long; green to golden-brown or reddish-tinged. Leaves clearly bilobed to 0.2 x the length of the leaf. Underleaves distinct along the entire stem. Median cells of leaf 18-30(32) x 22-40 |im; marginal c e l l s 12-24 x 16-26 p.m. Oil-bodies (1)2(3,4) per c e l l ; homogeneous, becoming segmented; spherical, varying from 3 -9 M-m i n diameter, or elongate and 4-8 x 6.0-17-5 M-m. Dioicous. Sporophytes infrequent, occurring i n August and September. Distribution: North Pacific; i n the study area, S t r i c t l y Subalpine-Alpine. New to North America; previously known only from Japan. Ecology: Growing on s o i l , humus, and gravel on streamlet banks or on slopes, or with l i t t e r and s o i l over boulders and outcrops; always i n open subalpine meadows. Substrate moist to wet, with seepage and snow-melt water. The species i s usually abundant where found. References: Amakawa 1959-NARDIA SCALARIS S. Gray Plants small to medium-sized, ca. 1.0-2.5 mm wide, clear lig h t green to red-tinged. Leaves orbicular to shortly reniform; apices entire. Underleaves distinct throughout the stem. Median c e l l s 22-32 x 28-40 (am. Oil-bodies (1)2-4(5) per c e l l ; homogeneous, shining, occasionally becoming segmented; spherical and (6)7-10(11) [im i n diameter, or elongate and (5)7-9(10) x (7.5)l0.0-17-5(-22.5) \m. 146 Dioicous. Plants seen were st e r i l e . Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia (N. scalaris s. l a t . ) ; the subsp. scalaris, to which plants from southwestern B r i t i s h Columbia belong, i s Disjunct North America — Western Eurasia; i n the study area, Suboceanic. Ecology: Predominantly occurring i n humid locations at low elevations where commonly growing on sandy s o i l (or over rock) on banks along streams, t r a i l s , and roads. Substrate mesic to moist, occasionally wet with seepage. The species also occurs infrequently at subalpine elevations, on s o i l or humus on streambanks, or on rock walls and outcrops. Here the substrates are moist to wet, with seepage or snow-melt water, and open. Subfamily MYLIOIDEAE MYLIA S. Gray Plants with rhizoids scattered along ventral stem. Leaves orbicular to ovate, succubously inserted, entire. Median leaf c e l l s very large (45-55 um long). Trigones very large and bulging. Underleaves lanceolate. Asexual reproduction by 2-celled, fasciculate gemmae. References: Inoue and Yang 1966. 1. Cuticle smooth; oil-bodies globular, appearing l i k e grape-clusters M. ANOMALA 1. Cuticle formed of angular plates, extremely coarse; oil-bodies granular . M. TAYLORI MYLIA ANOMALA (Hook.) S. Gray Plants large, 2-3 mm wide, translucent yellowish-brown to yellowish-green. Leaves orbicular-ovate. Median leaf cells very large, ca. to 147 4-5-55 fim. Trigones large, bulging. Cuticle smooth. Oil-bodies varying from 3_18 per c e l l , consisting of large globules or grape-clusters; spherical and 5-10 |im i n diameter, or elongate and 7.5-10.0 x 12.5-20.0 \xm. Gemmae 2-celled, greenish, spherical to ovoid, produced on the apices of (upper) lanceolate leaves. Dioicous. Plants from the study area were st e r i l e . Distribution: Circumboreal - Missing i n Eastern Asia, (occurring i n peat bogs); i n the study area, Widespread (elevation-independent, i n peat bogs). Ecology: A characteristic species of Sphagnum bogs, where i t occurs on wet humus and peat i n open sites. MYLIA TAYLORI (Hook.) S. Gray Plants large to robust, 3 _5 mm wide; usually b r i l l i a n t red to purplish-red. Leaves orbicular-ovate, entire. Median c e l l s very large, 46-60 x 60-80 (im. Trigones coarse, angular to nodulose. Cuticle extremely coarse, formed of heavy plates. Oil-bodies 10-18 per c e l l , granular, ovoid-elongate and 7.5-12.5 x 10.0-17.5 (JJtn. Dioicous. Plants from the study area were s t e r i l e . Distribution: Circumboreal; i n the study area, S t r i c t l y Oceanic. Ecology: Occurring i n very humid forests or bogs near the ocean, where i t grows on decaying logs, on tree trunks (red cedar), and on humus or peat on the ground. Substrate mesic to moist, partially shaded to open. Family GYROTHYRACEAE GYR0THYRA Howe 148 GYROTHYRA UNDERWOODIANA Howe Plants large to robust, 2-4 mm wide; light green with reddish to purplish secondary coloration. Rhizoids originating from dark purple pads on the ventral surface of the stem (these pads infrequently lack the distinctive purple coloration). Leaves orbicular to ovoid, entire or emarginate. Median leaf cells (24,28)34-40 x 43-68 |im; marginal cells ± forming a border, with one row of cells appearing larger than those immediately inside, and becoming thick-walled; basal cells larger than the median cells. Trigones bulging. Cuticle smooth. Oil-bodies 2-6 per cell, colorless to yellowish, granular; ovoid-ellipsoid, 5.0-6.5 x 7.5-17.5 M-™--Gemmae occasionally produced at stem apices, unicellular, spherical, green. Dioicous. Sporophytes found in April; elongate capsules with spiral dehiscence. Distribution: Endemic to Western North America; in the study area, Suboceanic. Ecology: Typically growing on sandy or clayey soil, in disturbed sites, e.g., roads, ditches, roadside banks, over uprooted trees, or along streams. Substrate mesic to moist, open. The species is usually abundant. References: Howe 1897a; Schuster 1955a, 1972b. Family GYMNOMITRIACEAE References: Muller 1942a, Kitagawa 1963a. GYMN0MITRI0N Corda Plants minute to small, shoots ca. 250 |im wide below, to 750 |im wide near apex; pale whitish-green to light golden-brown; typically growing in compact mats on exposed rock. Leaves bilobed, transversely inserted, 149 densely imbricate and appressed. Underleaves absent. Asexual reproduction lacking. Dioicous. Perianths absent. Perigynium weak (calyptral perigynium present). References: Kitagawa 1959• 1. Plants brownish-red to blackish (See Marsupella) 1. Plants pale whitish or silvery-green to light golden-brown . . . . 2 2. Sinus of leaf lunate, crescent-shaped; (plants becoming warm chestnut-brown) (See Marsupella condensata) 2. Sinus of leaf acute 3 3. Apices of leaf lobes acute; leaf sinus open but sharply acute; margins of leaf lobes plane, or becoming somewhat appressed or curved toward the next leaf upward on the stem; plants occurring at subalpine elevations G. GONGINNATUM 3. Apices of leaf lobes rounded; leaf sinus acute, "pinched" closed (by leaf lobes); margins of leaf lobes often becoming revolute; plants occurring at both low and subalpine elevations G. OBTUSUM GYMNQMITRION GONGINNATUM (Lightf.) Gorda (See above key and generic description.) Sporophytes occasional, found from August through October. Distribution: Circumboreal, Arctic-Alpine, Bipolar; in the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on perpendicular to slanting faces of bare rock (occasionally with i t s own humus or some accumulated s o i l ) , on boulders, outcrops, and c l i f f walls i n exposed sites. Substrate mesic to moist with seepage and snow-melt, but subject to drying out. 150 GYMNOMITRION OBTUSUM (Lindb.) Pears. (See above key and generic description.) Sporophytes occasional, May through October. Distribution: Disjunct Western North America — Greenland — Western Eurasia; i n the study area, Widespread (elevation-independent). Ecology: Growing on exposed rock (occasionally with some sand or s o i l ) , on boulders, outcrops, and c l i f f s , i n humid locations on boulder-slopes, i n forests, and on ocean beaches. Substrate mesic, becoming moist with seepage, or dry; usually open. MARSUPELLA. Dum. Plants with leaves bilobed, transversely inserted, usually at least somewhat erect-spreading. Underleaves absent. Oil-bodies typically 2 per c e l l . Asexual reproduction lacking. Perianth small, immersed, borne on a tubular perigynium. No bracteole. References: Kitagawa i960. 1. Leaves broader than the stem; plants relatively common i n southwestern British Columbia 2 1. Leaves narrower (or barely broader) than the stem; plants rare i n southwestern B r i t i s h Columbia 5 2. Plants minute, ca. 36O jam wide, less than 1 cm high; growing i n dark reddish-brown to blackish mats on exposed alpine ground M. BREVISSIMA 2. Plants larger; habitat various 3 3. Plants paroicous; purplish-brown, appearing scorched; leaf apices acute M. SPARSIFOLIA 3. Plants dioicous; color various, but i f purplish-brown to scorched, then not with acute leaf-lobe apices 4 151 4. Plants green or with red pigments, never appearing scorched; with the tendency for leaves to stand sti f f l y away from the stem; usually with some leaves having revolute margins; apices of leaf lobes usually ± acute; cortical cells of stem thick-walled; common M. EMARGINATA 4. Plants dull green to purplish-brown to scorched and blackish, lacking red pigments; usually tending to be very soft and lax in texture; apices of leaf lobes usually very broadly rounded; stem cortex a hyalodermis; usually in subalpine meadows . . . . M. SPHACELATA 5. Plants julaceous, Gymnomitrion-like. warm chestnut-brown; leaf sinus lunate; Garibaldi region, subalpine . . . . M. GONDENSATA 5. Plants not julaceous, dull reddish-brown to blackish; leaf sinus obtuse to acute; hyperoceanic 6 6. Plants ± erect, growing in crowded patches; lobe apices and sinus of leaves obtuse to narrowly rounded M. ADUSTA 6. Plants stringy, Gephaloziella-like, forming intricate mats; lobe apices and sinus of leaves very acute M. STABLER! MARSUPELLA ADUSTA (Nees emend. Limpr.) Spruce Plants minute to small, less than 400 \m wide in lower portions, to 1 mm wide below shoot apices; shoots erect, only J-k mm high; brilliant red (with transmitted light) to golden-red to red-brown, appearing blackish when dry; growing in dense, crowded patches. Stems, in transverse section, 10 cells in diameter; the cells a l l ± similar in size (with measurements similar to those for median leaf cells); cortical cells golden, medullary cells hyaline; a l l cells with large, distinct trigones. Shoots frequently branched; lower portions of stem with flagelliform branches. Rhizoids not seen. Leaves transversely inserted, concave, ovate-rectangular, longer than wide, the length 1.1-1.5 x "the width; on upper portions of shoots ca. 320-430 x 480 |_im, leaves on lower portions of shoots smaller; leaf sinus descending 0.2-0.3 x the leaf length, obtuse to narrowly rounded; lobe apices obtuse to narrowly rounded. Walls of leaf cells golden, though 152 l i t t l e distinguishable from trigones, which are very large, confluent, irregular to rectangular, appearing to occupy approximately as much of the leaf-area as do the c e l l lumina. Oil-bodies 2(-3-<4) per c e l l , spherical, |im i n diameter, very fa i n t l y papillose. Sex organs not seen; sporophytes lacking on plants from the area of this f l o r a . According to Kitagawa (1963a-), M. adusta i s monoicous (usually paroicous, occasionally autoicous); perianth lacking; wall of sporophyte capsule 2-stratose; nodular thickenings present on both inner and outer walls; elaters 9 UJH i n diameter, with J-k spirals; spores brown, 9 -(im i n diameter. N. Kitagawa (pers. comm.) believes that this plant may not be Marsupella adusta. Sporophyte-bearing material should be sought i n the future to aid i n clarifying the identity of this species. Distribution: Disjunct Western North America — Western Eurasia — Eastern Asia; i n the study area, Hypereceanic. Ecology: Growing on rock i n a dry, open meadowy area of an extensive bog near the Pacific Ocean on Brooks Peninsula. The species was abundant at the site. MARSUPELLA BREVISSIMA (Dum.) Grolle Plants minute, only ca. 3&0 (im wide, less than 1 cm high; dark reddish- or purplish-brown to fuscous, blackish. Rhizoids sparse, color-less or purple. Leaves small, concave but not tightly appressed. Apices of leaf lobes acute; sinus acute to rectangulate. Median leaf c e l l s 8-12 x 10-16 \m\ marginal c e l l s 8-10 urn long. Cell walls somewhat thickened. Cuticle smooth. Sexual condition apparently variable; according to Schuster ( 1 9 7 ^ ) , 153 normally paroicous, occasionally autoicous, sometimes with (pseudo?) dioicous shoots. Perianth absent or greatly reduced. Sporophytes found in the study area in August. Elaters 3-4-spiral. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (arctic-alpine); in the study area, Strictly Subalpine-Alpine. Ecology: Growing on soil or compacted humus (peaty soil) on the ground in alpine sites, especially in late-snow areas, occasionally over low, flat rock outcrops. Substrate mesic or moist to wet with snow-melt seepage; sites open and exposed. MARSUPELLA GONDENSATA (Angstr. in Hartm.) Schiffn. Plants minute to small, 280-560 \m wide; green to red-gold to chestnut-brown; threadlike, julaceous, resembling Gymnomitrion. Rhizoids lacking except below the gynoecia. Leaves subcontiguous to imbricate, quadrate-ovate, concave, 232-328 [im wide x 280-424 |im long; lobes incurved, acute; sinus lunate, very shallow, 0.05-0.10 x the leaf length. Median cells 14-20 x 19-24 |im; marginal cells 12-20 x 16-20 |im. Trigones small to larger and bulging. Cuticle smooth. Oil-bodies 2-3(4) per cell, granular; spherical and 5_8 |im in diameter, or elongate and 5-10 x 7-12 |im; occurring in a l l leaf cells, including marginal cells (to differentiate from Gymnomitrion apiculatum). Dioicous. Fertile shoots ± clavate. Gynoecial shoots larger than sterile shoots, especially toward the apex; leaves increasing in size toward the bracts. Perianth immersed; mouth cells fingerlike. Sporophytes not observed. Distribution: Disjunct Western North America — Greenland — 154 Western Eurasia, (arctic-alpine); i n the study area, S t r i c t l y Subalpine-Alpine (known only from the Garibaldi region). Schuster (1974: 111) stated that M. condensata i s considered "a rather rare arctic species, apparently very rare i n alpine situations south of the main body of the Tundra (and unknown from such situations i n North America)." At the time this plant was collected i n B r i t i s h Columbia, the species was unknown from continental North America. Very recently, however, M. condensata has been reported from Mt. McKinley i n Alaska (Vana 1976b). Ecology: Growing abundantly on s o i l and humus over boulders and rock outcrops on NE-facing slopes on the shore of Garibaldi Lake, elevation 1468 m. The substrate i s mesic to moist, open, and the habitat i s cold-alpine as a result of the proximity of this large lake, which i t s e l f i s fed by glaciers. A smaller population of M. condensata was found on the Diamond Head T r a i l , elevation ca. 1615 m, growing on moist s o i l over a c l i f f above subalpine meadows. MARSUPELLA EMARGINATA (Ehrh.) Dum. Plants medium-sized to large, ca. 2 mm wide; ligh t green to darker golden- to olive-green, with red pigments usually present; plants occasionally becoming red, never appearing scorched or blackish. Transverse section of stem showing 1 layer of thick-walled hyaline cortical cells; subcortical cells thick-walled, cream-colored to yellowish, smaller i n diameter than the cortical c e l l s ; and thin-walled medullary c e l l s whose diameters are ± equal to those of the corti c a l c e l l s . Rhizoids colorless or red. Leaves frequently standing s t i f f l y away from the stem, and some leaves on each plant with revolute margins. Leaf lobe apices usually acute to obtuse, occasionally becoming rounded; sinus rounded and open to obtusely rectangular or acute. Median leaf cells 10-24 x (16,17)20-34 (im; marginal cells 10-16 u.m; basal cells at leaf middle thick-walled, much elongated. Cuticle usually smooth, occasionally becoming striate. Oil-bodies usually 2 per cell, granular, oblong-ovate or spherical; 5*0-7.5 |im in diameter, or (2.5)3.5-7.5(9.0) x 5.0-12.5(-l6.0) u.m. Dioicous. Antheridial stalks biseriate. Perianths often present. Sporophytes found from March through June (low elevations), July through September (subalpine). Elaters bispiral, (6)8-10 jim in diameter; spores papillose, 10-14 |im in diameter. Distribution: Circumboreal (M. emarginata s. lat.; the subsp. emarginata, to which plants from southwestern British Columbia belong, is Circumboreal - Missing in Eastern Asia); in the study area, Widespread (elevation-independent). Ecology: Primarily occurring over rock substrates, i.e., on rocks, boulders, outcrops, or c l i f f faces (on bare rock or with sand, l i t t e r , soil, or humus); occasionally on soil banks. Habitats include humid forests, subalpine meadows, or near streams and lakes. Substrate mesic to wet, often with seepage; partially shaded to open. MARSUPELLA SPARSIFOLIA (Lindb.) Dum. Plants small, to 5 _7 mm high; blackish when dry, appearing warm chestnut brown with purplish tinges under transmitted light; in the field often appearing scorched; lacking discrete red pigments. Stems in transverse section showing thick-walled, often dark brown cortical cells; golden-brown subcortical cells transitional in size and appearance to the cells of the medulla, which are thin-walled and hyaline. Rhizoids purple and/or colorless. Leaf lobes acute'to obtuse; sinus acute or obtuse, descending (0.22)0.25-0.35(0.40) x the leaf length. Median leaf cells 14-20 x 20-30 jam, or somewhat larger and 16-28 x 22-40 p.m; marginal cells 156 (12)14-26(30) nm. long. Cuticle striate. Oil-bodies 2-3(4) per c e l l , granular, spherical and 4-6 \m i n diameter. Paroicous. Antheridial stalk 2-seriate. Perianth immersed. Sporophytes found i n June (low elevation) and August and September (subalpine). Capsule wall 2-3 c e l l layers thick. Elaters bi s p i r a l , (8)10-12 p.m i n diameter; spores finely papillose 10-12(15) [im i n diameter. Distribution: Disjunct North America — Western Eurasia, Arctic-Alpine, Bipolar; i n the study area, Subalpine-Descending. Ecology: Growing on humus or humusy s o i l over rock outcrop-slopes i n humid locations. Substrate moist to wet with seepage; generally open. MARSUPELLA SPHACELATA (Gies. i n Lindenb.) Dum. Plants medium-sized to large, 1.0-2.5 mm wide; du l l , clear green to blackish and appearing scorched; with transmitted light, appearing warm chestnut brown with purple tinges, discrete red pigments lacking; plants soft and lax i n texture. Transverse section of stem showing a hyalodermis of large, thin-walled hyaline c o r t i c a l cells whose diameters are approximately 2 times the diameters of the thin-walled medullary c e l l s ; subcortical c e l l s thick-walled, smaller i n diameter than the cortical c e l l s , frequently becoming brownish. Rhizoids colorless. Leaf lobes with apices almost always broadly rounded, occasionally obtuse; sinus acute. Median leaf c e l l s 12-20 x 22-30 [im; marginal c e l l s (10) 14-18 \m; basal ce l l s not thick-walled, not elongated. Cuticle striate. Dioicous. Perianths present. Sporophytes found i n August to September at subalpine elevations. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (montane); i n the study area, 157 Subalpine-Descending. Ecology: Predominantly subalpine, occurring i n open meadows, on s o i l or with i t s own humus on outcrops or rocks or on the ground i n streamlets, on banks, i n depressions, or i n pool margins; substrate moist to wet or submerged. Less frequently descending to low elevations and growing i n humid places with sand, humus, or s o i l over rock outcrops; substrate mesic to wet with seepage, p a r t i a l l y shaded to open. MARSUPELLA STABLERI Spruce ^ Plants minute to small, 200-500 UJH wide, 1-2 cm long, f i l i f o r m , Gephaloziella-like, brownish-red to dark red or blackish; growing i n dense, intricate, stringy mats or patches. Rhizoids few. Leaves remote to contiguous, small, ovate, narrower than the stem, imbricate, tightly appressed to the stem (at least along median and lower portions of the stem, where the leaves even become scalelike). Leaf lobes narrowly triangular-lanceolate, acute; sinus deeply V-shaped, descending to 0 . 2 5 -0.50 x the leaf length. Cells small, ca. (8 -) l0 -18 x 18-20 p.m. Trigones small to coarse but indistinct; c e l l walls at least slightly thickened. Cuticle smooth. Dioicous. F e r t i l e portions of both male and female shoots club-like, suddenly larger than the tightly appressed-leaved, f i l i f o r m , s t e r i l e portions of the shoots. Perianth slightly shorter than the female bracts. Bracteole absent. Sporophytes not found on plants from the study area, but elaters 2-spiral, 6-7 |im i n diameter; spores 15-18 |om i n diameter, according to Schuster (1974), who treats this plant as a variety of M. boeckii (Aust.) Lindb. Distribution: Disjunct North America — Western Eurasia; i n the study area, Hyperoceanic (but with a single population known from subalpine 158 meadows near Mt. Garibaldi; these plants were very recently identified by N. Kitagawa). Ecology: Growing very abundantly on s o i l over rocks. On Brooks Peninsula, pa r t i a l l y submerged i n standing water along a streamlet i n an extensive lodgepole pine bog. Family SCAPANIACEAE References: Muller 1905; Buch 1922, 1928; Amakawa and Hattori 1953, 1954, 1955; Amakawa 1964, 1968; Grolle 1965b. DIPLOPHYLLUM Dum. emend. Lindb. Plants small to medium i n size, 1.0-2.5 mm wide (subg. Diplophyllum), or large to robust, 4-5 mm wide (subg. Macrodiplophyllum). Leaves complicate-bilobed, dorsal lobe smaller, the lobes narrow and Ungulate, widely divergent, lobe margins denticulate. Leaf cells v i r t u a l l y noncollenchymatous i n subg. Diplophyllum, or with large trigones i n subg. Macrodiplophyllum. Gemmae often present. Perianth deeply plicate. References: Amakawa and Hattori 1955-1. Plants large to robust, 4-5 mm wide; leaves decurrent ventrally; leaf c e l l s strongly collenchymatous; gemmae ± (rounded to) cubic, J-k-celled. i n part D. (MACRODIPLOPHYLLUM) PLICATUM 1. Plants small to medium-sized, 1.0-2.5 mm wide; leaves not decurrent ventrally; leaf cells thick-walled, not or barely collenchymatous; gemmae stellate, 1-2-celled . 2 (D. Subg. DIPLOPHYLLUM) 2. Leaf lobes with a distinct median band of elongated, whitish cells (vitta) D. (DIPLOPHYLLUM) ALBICANS 2. Leaf lobes lacking a distinct v i t t a 3 159 3. Plants shiny, yellow-green to reddish; median leaf cells of lower portion of ventral lobe elongate, to 30-60 UJH long; apical leaf c e l l s to ca. 10-12 \m long; gemmae lacking (in the literature, said to be rare, 1-celled); common on roadside s o i l banks and ditches . . . D. (DIPLOPHYLLUM) 0BTUSIF0LIUM 3. Plants dull olive or golden-green, lacking red pigments; median leaf c e l l s of ventral lobe shorter,- to 22-26 UJH long; apical leaf c e l l s to 8-10 ujn long; gemmae usually abundant, green, 2-celled; common on rock outcrops, especially at subalpine elevations D. (DIPLOPHYLLUM) TAXIFOLIUM DIPLOPHYLLUM (DIPLOPHYLLUM) ALBICANS (L.) Dum. Plants medium-sized, 1.5-2.5 mm wide; green to golden, with red pigments. Leaf lobes vittate, with a distinct median line formed by elongated whitish c e l l s . Gemmae frequent, stellate, 1-celled, yellow-green. Dioicous. Perianth mouth c i l i a t e . Sporophytes found i n A p r i l . Distribution: Circumboreal5 i n the study area, Widespread (essentially low-elevation). Ecology: Growing i n humid, moist sites on several types of organic or inorganic substrates, including decaying logs, peaty hummocks i n bogs, trunks of l i v i n g trees, rocks i n streams, c l i f f faces, outcrop crevices, streambanks, and roadside s o i l banks. Substrate mesic to wet, often with seepage or spray; pa r t i a l l y shaded to open. DIPLOPHYLLUM (DIPLOPHYLLUM) 0BTUSIF0LIUM (Hook.) Dum. Plants medium-sized, 1.5-2.5 mm wide; yellow- or golden-green, to golden-brown, or becoming reddish. Leaf lobes not vittate. Median cells of leaf 30-60 |jjn long. Gemmae not found; reported to be very rare, 1-celled, brownish (Schuster 1974). Paroicous. Perianths almost always present; the mouths with long teeth or c i l i a . Sporophytes found i n A p r i l , May, and October. 160 Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia; i n the study area, Widespread (essentially low-elevation). Ecology: Characteristically occurring on s o i l i n open, disturbed sites, as on roadside s o i l banks, with humus over rock outcrops, and over roots of trees. Substrate mesic, generally open. DIPLOPHYLLUM (MACRODIPLOPHYLLUM) PLICATUM Lindb. Plants resembling Diplophyllum subg. Diplophyllum, though larger to robust, ca. 4-5 mm wide at widest part of shoot, green to brownish. Stems becoming dark, appearing purplish-black; with transmitted li g h t , dark reddish-brown; sparingly branched. Cortical c e l l s of stem i n 2-4 layers, the walls thickened (transverse section). Rhizoids few. Leaves distant to contiguous, ± transversely inserted, complicate-bilobed, decurrent both dorsally and ventrally. Dorsal lobes appressed to the stem, spatulate, the length ca. 0.6-0.7 x the length of the ventral lobe; margins entire or somewhat denticulate toward the rounded apex. Keel 0.4 x the length of the ventral lobe. Ventral lobes widely spreading, ligulate, slightly falcate, narrower near base, the margins entire or somewhat denticulate toward the rounded apex. Marginal c e l l s of leaf 10-12 [im long, collenchymatous or becoming thick-walled; median cells 16-26 x 20-40 |_im, the walls thickened, trigones large to ± bulging; basal c e l l s more elongate, 20 x 40-80 pm (Amakawa and Hattori 1955)- Cuticle fa i n t l y papillose. Gemmae 2-4-celled, green to brownish, ovoid to rotund-cubic, 16-20 x 20-30 [im. Dioicous. Male inflorescence terminal or intercalary. Perianths cylindrical, not or l i t t l e compressed dorsiventrally, deeply plicate to the base; perianth-mouth 1/2 x or nearly as wide as the broadest part of the perianth, irregularly laciniate-fimbriate. Sporophytes not observed. 161 Persson (1949) elevated Macrodipiophyllum previously considered a subgenus of Diplophyllum, e.g., i n Buch (1928), to the generic level. In their revision of the Japanese Scapaniaceae, Amakawa and Hattori (1955) also treated Macrodiplophyllum as an independent genus. Schuster (1974: 197» 18l) discusses the features by which Macrodiplophyllum i s said to d i f f e r from Diplophyllum. These include: more robust size; 2-4-stratose stem cortex; ventral-intercalary branching; decurrent postical leaf base; larger, collenchymatous leaf cells; cubic, often 3 - ^ - c e l l e d gemmae; and longly fimbriate perianth mouth. He does not f e e l that these features are sufficient to establish Macrodiplophyllum as an independent genus, and so retains i t as a subgenus of Diplophyllum. However, Schuster (1974: 179, Footnote 3) states, "If the branching differences (ventral-intercalary i n Macrodiplophyllum, lateral-intercalary i n Diplophyllum) cited by Grolle . . . prove valid, i t may s t i l l be possible to retain Macrodiplophyllum as an independent genus." In plants from southwestern B r i t i s h Columbia, I found only lateral-intercalary branches, i.e., with branches originating near axils of (lateral) leaves. The presumptive difference i n branch origin i s therefore not a valid character and Macrodiplophyllum cannot be treated as an independent genus. Inoue (1974a) has recently placed Macrodiplophyllum as a subgenus of Diplophyllum• Brinkman (1940) briefly discusses a species he cal l s Diplophyllum  hyalinum Brinkman ms., based on material mixed i n with a species of Scapania collected by Macoun. He does not describe the "species," nor does he give a Latin diagnosis. Frye and Clark (1946) include Brinkman1s D. hyalinum i n their book; they add a Latin diagnosis, complete description, and il l u s t r a t i o n s . I have seen this material and regard i t as D. plicatum. Brinkman, however, separated his D. hyalinum from D. plicatum on the basis of variations in size and appearance of ce l l s at the bases of the leaves. 162 Distribution: North Pacific; i n the study area, Suboceanic. Ecology: On the west coast of Vancouver Island, growing on decaying logs, trunks of trees, and peaty rotting stumps, i n rainforest and Sphagnum bogs. Elsewhere, growing on sand or sandy humus over rocks (boulders, outcrops, rock walls) on stream banks and near waterfalls i n cold, humid places along cold mountain streams (at low elevations). DIPLOPHYLLUM (DIPLOPHYLLUM) TAXIFOLIUM (Wahlenb.) Dum. Plants medium-sized, 1-2 mm wide; du l l medium green to golden-green or olive. Median c e l l s to 22-26(30) UJII long; v i t t a lacking. Gemmae usually present and abundant, 2-celled, green. Dioicous. Perianths present on about half the specimens examined; perianth mouth hyaline, with short teeth. Sporophytes found i n June at low elevations, i n August through October at subalpine elevations. Distribution: Gircumboreal, Arctic-Alpine; i n the study area, Widespread (elevation-independent). Ecology: Growing on rock, i.e., often with humus, s o i l , or l i t t e r , over rocks, boulders, and outcrops; also growing on s o i l banks along t r a i l s and streams. Substrate generally mesic; open to partially shaded. DOUINIA Buch DOUINIA OVATA (Dicks.) Buch Plants medium-sized, 1-2 mm wide (to 2.5 mm wide at apices of female shoots); glaucous-opaque bluish-green to brownish. Leaves complicate-bilobed, the dorsal lobe smaller; apices of both lobes sharply acute. Underleaves absent. Gemmae not observed. Dioicous. Perianth oblong, deeply plicate to near base; the mouth broad, truncate, with long c i l i a . Sporophytes occasional, found i n 163 February, A p r i l through August, November (low elevations); June and July (subalpine). Distribution: Disjunct Western North America — Western Eurasia; i n the study area, Widespread (essentially low-elevation). Ecology: Occurring i n humid habitats; epiphytic on l i v i n g trees and shrubs i n forests and bogs, also growing on humus and with mosses over rock as on boulders on boulder-slopes, and on outcrops. Infrequent at subalpine elevations, where confined to the forest canopy (on mountain hemlock and yellow cedar). Substrate dry to mesic, usually partially shaded. SGAPANIA (Dum.) Dum. Plants with complicate-bilobed leaves, the dorsal lobe smaller; the lobes variously ovate, obovate, rectangular, orbicular, or reniform, oriented ± i n the same direction at an angle to the stem. Underleaves lacking. Leaf cells collenchymatous. Asexual reproduction by smooth, ovoid-ellipsoid gemmae. Dioicous. Perianths generally strongly flattened, the mouths broad and truncate. References: Muller 1905, 1942b; Evans 1907b, I936; Buch 1922, 1928; Amakawa and Hattori 1953, 1954, 1955; Amakawa 1964, 1968. Of the 13 species of Scapania now known from southwestern British Columbia, 12 belong to the subgenus Scapania. Scapania gymnostomophila, however, belongs to subgenus Kaalaasia Buch emend. Schust. Plants of subg. Kaalaasia possess only 1 (S. gymnostomophila) or 2-5 very large (nearly f i l l i n g c e l l lumen), persistent oil-bodies per c e l l , whereas oil-bodies of subgenus Scapania are relatively smaller and more numerous, varying from 2 164 to 15 per c e l l . The perianth i n subg. Kaalaasia i s contracted to the mouth, strongly plicate, and only weakly dorsiventrally compressed. This contrasts with the perianth of subg. Scapania, which i s strongly dorsiventrally compressed, with the mouth truncate and nonplicate. 2. Ventral lobes broadly cordate to reniform; keel strongly semicircularly arched, very short, i t s length less than 0.25 x the length of the ventral lobe; plants rare i n southwestern B r i t i s h Columbia, occurring i n Sphagnum bogs S. FALUDICOLA 2. Ventral lobes narrow, oblong-obovate, the width equal to 0.50-0.75 x the length; keel straight, i t s length ca. O.3-O.5 x the length of the ventral lobe 3 3. Leaf margins with coarse teeth; nongemmiparous lobes of leaves bordered, i. e . , with 2-3 rows of c e l l s along the margins equally thick-walled, noncollenchymatous; dorsal lobe of leaf with a free, c i l i a t e d extension crossing the stem; plants large to robust, 2.0-4.5 mm wide, very common on decaying logs at low elevations S. BOLANDERI 3. Leaf margins entire; nongemmiparous lobes of leaves collenchymatous, not bordered; dorsal lobe of leaf lacking a free, c i l i a t e d extension; plants small, 1.0-2.5 mm wide, generally growing on rock 4 4. Oil-bodies very large, brownish, persistent, mostly occurring only 1 per c e l l ; gemmae brown S. GYMN0ST0M0PHILA 4. Oil-bodies smaller, several per c e l l ; gemmae green to reddish . . . 5 5. Plants green or olive-green to golden-brown; margins of leaf lobes plane; perianth mouth densely c i l i a t e . . . . . . S. MUCRONATA 1. Leaves not decurrent ventrally 2 1. Leaves decurrent ventrally 6 5. Plants green to golden-brown with purplish-red pigments; apical margins of ventral lobes of leaves often becoming inrolled; perianth mouth entire or sinuate S. SCANDICA 165 6. Leaf cuticle extremely verrucose, with large, hemispherical papillae; (dorsal lobes subequal to the ventral lobes i n size; gemmae brown) SCAPANIA (new species) 6. Leaf cuticle smooth to moderately papillose, but never with large, hemispherical papillae 7 7. Ventral lobes narrow, 0.55-0.75 x a s wide as long; dorsal lobes transverse; plants small, 1-2 mm wide S. UMBROSA 7. Ventral lobes ± ovate, 0.7-1.0 or more tines as broad as long; dorsal lobes transverse or decurrent; plants medium to large or robust, 2.0-4.5 mm wide 8 8. Dorsal lobes decurrent 9 8. Dorsal lobes transverse 11 9. Keel strongly semicircularly arched, very short, i t s length no more than 0.15 x "the length of the ventral lobe S. PALUDOSA 9. Keel straight or only weakly arched, longer, 0.2-0.3 x the length of the ventral lobe 10 10. Leaf lobes not or weakly convex; leaf margins strongly, closely dentate-ciliate, the margins thus often appearing to have a whitish fringe; leaf c e l l s with trigones concave to bulging; perianth mouth c i l i a t e ; plants green to golden-brown, with red pigments; common on rock at low elevations S. AMERICANA 10. Leaf lobes strongly convex; leaf margins entire; leaf c e l l s with minute trigones or trigones lacking; perianth mouth truncate, entire; plants becoming blackish, "scorched;" infrequent, i n subalpine meadows S. ULIGINOSA 11. Leaf lobes of different sizes, the dorsal lobe ca. 0.6 x as long as the ventral lobe; keel 0.25-0.30 x the length of the ventral lobe S. UNDULATA 11. Leaf lobes subequal, the dorsal lobe 0.75-0.90 x as long as the ventral lobe; keel 0.50-0.65 x the length of the ventral lobe (rather lax, the plants thus appearing ruffly) 12 166 12. Plants lig h t golden-green, becoming deep reddish- to fuscous-brown; leaf margins entire; marginal c e l l s collenchymatous, i.e., the leaves not bordered; plants rare to infrequent, subalpine S. OBSGURA 12. Plants lig h t green; leaf margins dentate; marginal cells evenly thick-walled, i.e., the leaves bordered; plants relatively common, subalpine or descending to lower elevations along cold streams S. SUBALPINA SGAPANIA AMERICANA K. Mull. Plants medium to large, 2.0-3-5 mm wide; li g h t golden-green, becoming golden-brown or tinged with red pigments. Stem dark red-brown. Leaf margins finely fringed with regular, sharp teeth or c i l i a , 2-5 c e l l s long; margins often appearing bordered. Dorsal lobes decurrent, reniform; apex rounded; the length equal to 0.4-0.6 x the length of the ventral lobe. Ventral lobes decurrent, obovate, convex; apex rounded; the width 0.6-0.9 x the length. Keel somewhat arched; length ca. 0.3 x the length of the ventral lobe. Trigones concave to bulging. Cuticle papillose. Gemmae 2-celled, green to golden or red. Dioicous. Perianth mouth with long c i l i a . Sporophytes found i n March, April, August. Distribution; Endemic to Western North America; i n the study area, Widespread (essentially low-elevation). Ecology: Generally growing on rock (on bare rock or on i t s own humus or accumulated s o i l , on boulders, outcrops, and c l i f f s ) . Substrate dry to mesic, or becoming moist with seepage; open or partially shaded. References: Evans 1930, Amakawa'I967. SGAPANIA B0LANDERI Aust. Plants usually medium to large, 2.0-4.5 mm wide (occasionally smaller and then superficially resembling S. umbrosa, a species which does, however, 167 have red pigments); pure to olive-green, d u l l , becoming (golden) to brownish, lacking any red or vinaceous pigmentation. Stems dark reddish-brown. Leaf margins with coarse, large teeth, 2-3 c e l l s long with a broad base 2 c e l l s wide; marginal c e l l s appearing bordered (or occasionally vague). Dorsal lobes transverse, with a c i l i a t e d , free extension crossing the (dorsal) stem; reniform; apex rounded; the length 0.75 x the length of the ventral lobe. Ventral lobes not decurrent beyond the keel insertion; oblong, ± convex, the apex rounded; the width 0.6 x the length. Keel straight, the length 0.25-0.30 x the length of the ventral lobe. Trigones large, irregular to bulging and confluent. Cuticle rough-papillose. Gemmae 2-celled, green. Dioicous. Perianth mouth with long teeth or c i l i a . Sporophytes found from March through July, November. Distribution: North Pacific; i n the study area, Widespread (essentially low-elevation; subalpine occurrences confined to forest canopy). Ecology: Very common on decaying logs and stumps, i n mesic to moist, pa r t i a l l y shaded sites i n the forest. In very humid sites, also growing on rocks, boulders, and outcrops; substrate moist, partially shaded. Occasionally epiphytic on tree trunks, usually toward the base when at low elevations.' References: Evans 1930, Amakawa I967. SCAPANIA GYMN0ST0M0PHILA Kaal. Plants small to medium-sized, 1.6-2.3 mm wide; opaque, yellowish-green to blackish and somewhat "scorched." Stems green to brownish; i n transverse section, the cortex 1-stratose, consisting of small c e l l s . Leaf margins entire; marginal c e l l s not forming a border. Dorsal lobes 168 transverse, very small, ovate; apex acute; length 0.20-0.35 x the length of the ventral lobe. Ventral lobes not decurrent, narrow, oblong; apex acute; the width 0.5 x the length. Keel weakly arched, the length 0.5 x the length of the ventral lobe. Trigones lacking or small and concave. Cuticle faintly papillose. Oil-bodies 1 per c e l l i n most ce l l s , brownish, finely granular, very large and almost f i l l i n g the c e l l lumen. Gemmae 2-celled, brown. Dioicous. Plants from the study area were s t e r i l e . Perianths, according to the literature, plicate, weakly flattened, narrowed to the mouth, which i s c i l i a t e . Scapania gymnostomophila i s the only species of the genus treated i n this f l o r a which does not belong to subg. Scapania. I t belongs instead to subg. Kaalaasia Buch emend. Schust. This species was previously reported from B r i t i s h Columbia by Conklin (1925), based on a specimen collected by F. A. MacFadden; this report i s considered doubtful by Schuster (197*0. The species was also included, as Diplophyllum gymnostomophilum Kaal., on Brinkman's (193*+) l i s t of hepatics from the Pacific Coast. Brinkman (1937) subsequently found the specimen had been misidentified, and removed the species from his l i s t . Distribution: Circumboreal - Missing i n Eastern Asia, Arctic-Alpine, (calcareous substrates); i n the study area, Rare or Restricted (known only from limestone c l i f f s , Chilliwack River Valley). New to B r i t i s h Columbia. Ecology: Growing on s o i l and rock, base of limestone c l i f f . Substrate moist, open to pa r t i a l l y shaded. References: Evans 1936. 169 SGAPANIA MUGRONATA Buch Plants small, 1-2 mm wide, to 12 mm long; green to olive or golden-brown, lacking red pigments. Stem golden-green to brownish. Margins of nongemmiparous leaves entire; not bordered, the marginal cells collenchymatous. Dorsal lobes transversely inserted, narrowly ovate-rectangular; apex obtuse to acute or ± apiculate; the length 0.5-0.6 x the length of the ventral lobe. Ventral lobes not decurrent, narrowly oblong-ovate; apex obtuse or acute; the width 0.5 x the length. Keel length 0.44 x the length of the ventral lobe. Trigones distinct to moderately large. Gemmae 2-celled, green to reddish. Dioicous. Perianth mouth rather densely c i l i a t e . Sporophytes found i n June. Distribution: Circumboreal; i n the study area, Dry Northern-Interior. Ecology: Growing on s o i l over rock, on outcrops and boulder-slopes. Substrate mesic, exposed. References: Evans 1936. SGAPANIA OBSCURA (H. Arn. et C. Jens.) Schiffn. Plants medium-sized, 2.0-2.5 mm wide, light golden- (or brownish-) green to deep, r i c h reddish-brown, with both brown and red pigments. Stems the same color as the rest of the plants; i n transverse section, the cortic a l cells golden-brown, smaller i n diameter than the hyaline cells of the medulla, which are ± abruptly larger. Leaves lax, ruffly; margins entire, not bordered, the marginal c e l l s collenchymatous. Dorsal lobes transversely inserted, rounded-quadrate to rounded-oblong, extending across and slightly beyond the stem, the length equal to O.65-O.8O x that of the ventral lobes, apex blunt to rounded. Ventral lobes l i t t l e larger than dorsal, ± ovate, the apex blunt to rounded, the width 0.6-1.0 x the length. 170 Keel ± straight to weakly arched, ca. O.5O-O.65 x the length of the ventral lobe. Median c e l l s of leaf ca. 20 x 30-35 P-m. Trigones small. Cuticle weakly papillose. Gemmae 1-2-celled i n the Bri t i s h Columbia plants, light orange-red to bright purplish-red, 10-12 x 14-22 |im. Dioicous. Perianth and sporophyte unknown (Schuster 1974). Scapania obscura was f i r s t identified among B r i t i s h Columbia hepatics by S. Hattori (W. B. Schofield, pers. comm.; WBS 35423. 35425, 35452). Scapania obscura i s very similar to S. subalpina, and i s separated from i t by: l) lack, of a border along leaf margins; 2) entire leaf margins, with no trace of teeth; and 3) presence of brown and red pigments (Schuster 1974, Schuster and Damsholt 1974). Schuster (1974: 547) states, "I remain sceptical of S. obscura as a species and suspect that, under very exceptional environmental conditions, leaf border formation may be suppressed i n S. subalpina, even though collenchyma may form." Distribution: Disjunct North America — Western Eurasia, (arctic-alpine); i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on humusy s o i l on a subalpine meadow slope, elevation over 1800 m. Substrate moist to wet, with seepage from melting snow; open. The species was abundant at the site. SCAPANIA PALUDICOLA Loeske et K. Mull, i n K. Mull. Plants large to robust, 2.5-4.0 mm wide; pale, transparent yellow-green to golden. Stem becoming golden to golden-brown ventrally. Leaf margins entire to emarginate; marginal c e l l s collenchymatous. Dorsal lobes decurrent, reniform to heart-shaped; apex acute (apical c e l l may form a tooth of up to 3 c e l l s ) ; the length 0.5 to 0.7 x the length of the ventral lobe. Ventral lobes not decurrent, orbicular to heart-shaped; apex acute; 171 the width equal or greater than the length. Keel highly arched, semicircular, very short; i t s length 0.1 x the length of the ventral lobe. Trigones bulging, sinuate. Cuticle striate. Gemmae 2-celled, (green to) brownish-red. Dioicous. Perianth mouth somewhat distantly c i l i a t e ; c i l i a consisting of 1-4 quadrate c e l l s i n a row. Sporophytes not found. Distribution: Circumboreal (in peat bogs); i n the study area, Rare or Restricted (known only from Joffre Creek). New to Brit i s h Columbia. Ecology: Growing over wet humus and Sphagnum i n an open bog. References: Evans 1916. SGAPANIA PALUDOSA (K. Mull.) K. Mull. Plants large to robust, 2.5-4.5 mm wide; transparent or translucent, pale pure green to yellow-green, to golden-brown or purplish-red-brown (these dark red forms appearing blackish when dry). Stems dark red-brown, contrasting greatly with pale color of the leaves i n lighter-colored forms. Leaf margins with teeth, or becoming entire, especially below on lower parts of leaves or plants; bordered. Dorsal lobes decurrent, reniform to heart-shaped; apex rounded-obtuse; length O.65 x the length of the ventral lobe. Ventral lobes decurrent, orbicular to ovate, the apex rounded; width ± equal to the length. Keel highly arched, very short; 0.15 or less x the length of the ventral lobe. Trigones small or absent, and c e l l walls thin; cuticle smooth. Gemmae lacking. Dioicous. Perianth mouth truncate, denticulate to ± entire. Sporophytes not observed. Distribution: Circumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. 172 Ecology: Hygrophytic, growing on rocks, rock outcrops, or on s o i l , humus, and l i t t e r on the ground; occurring i n subalpine meadows and i n and adjacent to subalpine streams; also descending to low elevations, where found on rock outcrops, c l i f f s , roadside banks and ditches, along cold streams, and i n west coast bogs ( a l l i n cold-humid locations). Substrate always wet, usually open. SCAPANIA SCANPICA (H. Arn. et Buch) Macv. Plants small to medium-sized, 1-2 mm wide; green to golden-brown, with red or vinaceous pigments i n c e l l walls and occasionally spreading to leaf bases and male bracts. Stems golden- to red-brown. Leaf margins entire except where gemmae are produced, then often becoming irregularly dissected, ragged; margins collenchymatous, not bordered. Dorsal lobes transversely inserted, triangular-ovate; the apex rounded, obtuse, or acute; length 0.7-0.8 x the length of the ventral lobe. Ventral lobes not decurrent, Ungulate to narrowly ovate, narrow, the width 0.50-0.75 x "the length, rather "sturdy" i n appearance; apex broadly rounded, margins becoming inrolled. Keel length 0.33-0.55 x the length of the ventral lobe. Gemmae 2-celled, green to reddish. Diocious. Perianth mouth entire or sinuate (occasionally with one or more isolated, small, smooth protuberances). Sporophytes found i n June and August. Distribution: Disjunct North America — Western Eurasia; i n the study area, Subalpine-Descending. Ecology: Usually growing on sandy s o i l (occasionally with humus or among bryophytes and lichens) over rock substrates, on boulders on boulder-slopes, on rock outcrops, and on c l i f f faces, i n ± exposed, dry to mesic sites; infrequent to abundant. Occasionally growing on decaying 173 wood, as on sand over a decaying stump near Lindeman Lake, and on a decaying log by Joffre Greek. Schuster (1974: 44-9) states that while Buch (1928) mentions wood as a substrate for this species, he himself has collected S„ scandica on wood only once, i n Scotland. SGAPANIA SUBALPINA (Nees i n Lindenb.) Dum. Plants medium to large, 2-4 mm wide; light green to golden-green. Stem green to (purplish-)brown. Leaf margins with minute teeth, bordered. Dorsal lobes transverse, ovate, the apex rounded; almost equal i n size to the ventral lobe, the length 0.75-0.90 x the length of the ventral lobe. Ventral lobes decurrent, ovate-obovate5 apex rounded; width 0.7-1.0 x the length. Keel O.5O-O.65 x the length of the ventral lobe. Trigones lacking or small. Cuticle smooth. Gemmae 2-celled, green to reddish. Dioicous. Perianth mouth with minute teeth. Distribution: Circumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. Ecology: Growing primarily i n subalpine meadows, and also descending along cold streams into the subalpine forest, occasionally reaching low elevations. Characteristically growing on sand, often over rocks or decaying logs, on banks or i n streams and rivers. Substrate moist to wet, subject to inundation; open to par t i a l l y shaded. SCAPANIA ULIGINOSA (Sw. ex Lindenb.) Dum. Plants large to robust, 3-4 mm wide; golden- to olive-green, to vinaceous or purplish-brown, appearing blackish and scorched. Stems dark, reddish-brown. Leaf margins entire. Dorsal lobes decurrent, broadly ovate to semicircular, convex; apex broadly rounded; length 0.7 x the length of the ventral lobe. Ventral lobes decurrent, broadly obovate, 174 strongly convex; apex broadly rounded; width approximately equal to the length. Keel ca. 0.2 x the length of the ventral lobe; only slightly arched. Trigones lacking or minute; c e l l walls becoming somewhat thickened. Cuticle ± smooth. Gemmae lacking. Dioicous. Perianths not observed; according to Schuster (1974), truncate, the mouth entire. Distribution: Circumboreal - Missing i n Eastern Asia, Arctic-Alpine; i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Hygro- to hydrophytic, growing on sand, gravel, or over rock outcrops i n open subalpine meadows or parkland. Relatively rare. SCAPANIA UMBROSA (Schrad.) Dum. Plants small to medium-sized, 1-2 mm wide; light, bright green to golden to red. Stems becoming dark reddish-brown. Leaf margins with coarse, broad-based teeth; not bordered. Dorsal lobes transverse, heart-shaped; apex acute; length 0.5 x the length of the ventral lobe. Ventral lobes decurrent, narrowly obovate, the apex acute; width 0.55-0.75 x "the length. Keel 0.33 x the length of the ventral lobe. Trigones concave to bulging and confluent. Cuticle papillose. Gemmae 2-celled, green to red. Dioicous. Perianth mouth entire, sinuate. Sporophytes occurring i n Apri l through June; found i n late August on an infrequent subalpine plant. Distribution: Disjunct North America — Western Eurasia; i n the study area, Widespread (essentially low-elevation). Ecology: Occurring i n humid forests, primarily on decaying logs or stumps. Substrate mesic to moist, partially shaded to open. SCAPANIA UNDULATA (L.) Dum. Plants large to robust, 2-4 mm wide; clear green, frequently with 175 leaf bases or entire plants becoming dark reddish-purple, appearing blackish. Stem dark red-brown to blackish. Leaf margins with even teeth consisting of 1-2 cells, these teeth varying from large to minute or vestigial; marginal cells forming a border. Dorsal lobes transverse, ± reniform; apex ± rounded (to acute with erosion from gemmae production); length 0.6 x the length of the ventral lobe. Ventral lobes decurrent, round (to ovate); apex rounded; width 0.80-0.95 x the length. Keel 0.25-0.30 x the length of the ventral lobe. Trigones lacking and cell walls ± even, or with small trigones. Cuticle faintly to distinctly papillose. Gemmae 2-celled, green to reddish. Dioicous. Perianth mouth dentate. Distribution: Circumboreal; in the study area, Widespread (elevation-independent). Ecology: Always associated with seepage or flowing water; growing on sand, s i l t , and rock, or on decaying logs on streambariks, in streams and. rivers, and on roadside banks and ditches. Substrate moist to wet, often with spray or submerged; open or partially shaded. References: Glime 1970.. SGAPANIA (new species) Plants small, 880-1280 |im wide, 9-15 mm long; dark brownish-green, resembling Marsupella in habit. Stems dark reddish-brown. Leaves distant to subimbricate; the lobes subequal, squarrose;.margins entire or with a few minute, widely scattered, 1-celled teeth; marginal cells thin-walled, collenchymatous. Dorsal lobes transverse to very shortly decurrent, ovate; the apex obtuse to acute; antical base broadly rounded, extending across and beyond the stem; length 0.70-0.86(0.94) x the length of the ventral lobe. Ventral lobes long-decurrent, ovate, reflexed; apex obtuse to acute; 176 the width 0.5-0.7 x the length. Keel straight or weakly arched, i t s length ca. 0.4-0.6 x the length of the ventral lobe. Median ce l l s 16-20 x 16-24(30) p;m; marginal c e l l s 9-12 x 10-l4(l6) |im. Trigones small to bulging and occasionally confluent. Cuticle very coarsely verrucose, with 3-8 relatively huge, hemispherical papillae per c e l l . Gemmae 2-celled, reddish- to golden-brown, spherical to ovoid, 10-13 (im i n diameter, or 12-16.x 14-20(24) jam; forming dark brown masses at stem apices. Sex organs, perianth, and sporophytes unknown. This species, which belongs i n Subgenus Scapania, Sectio Aequilobae. K. Mull, emend. Buch, w i l l be f u l l y described and ill u s t r a t e d i n a future manuscript. Distribution: Endemic to Western North America; i n the study area, Subalpine-Descending. Ecology: Growing on thin mats of i t s own humus on clayey s i l t over rock i n fast-flowing, glacier-fed, mountain streams, i n subalpine forest (elevation 1432 m), and at a lower elevation (ca. 518 m) by a waterfall. Substrate moist to wet or submerged, depending on the seasonal water-level; open to par t i a l l y shaded. Family ANTHELIACEAE ANTHELIA (Dum.) Dum. Plants minute, less than 500 p.m wide, moss-like, crawling, essentially green to brownish or blackish, but this color often becomes obscured by a "mouldy-looking" whitish, crystalline substance, thus making the plants appear grayish to whitish. Branches frequent, generally lateral-terminal. Leaves ± transversely inserted, closely imbricate, longer than broad; 177 deeply divided into two long, narrow, acute lobes. Underleaves vi r t u a l l y identical to leaves. Leaf c e l l s equally thick-walled. Oil-bodies absent. Asexual reproduction lacking. References: Schuster 1964b. 1. Plants relatively robust, 8-35 mm long, julaceous, crawling, dark green to blackish or becoming "mouldy;" dioicous; perianth oblong, relatively long-emergent; elaters with 2 ( -3) obviously broad ( 2 - 4 |im) spirals; plants occurring at both low and subalpine elevations A. JULAGEA 1. Plants minute, less than 10 mm high, erect when crowded, forming white crusts; paroicous; perianth ovate, ± enclosed by bracts; elaters with 2 -3 narrow (2 jam broad or less) spirals; always occurring at subalpine or alpine elevations . . A. JURATZKANA The primary characteristics used to separate the species are given i n the above key. While the most minute crustose forms of this genus usually seem safely referable to A. juratzkana, and the most robust, dark green to blackish crawling forms, to A. julacea (especially at low elevations), there are many intermediate forms, and size alone cannot be used for determination of species. The shape of the perianth and i t s degree of emergence from the bracts seem more reliable but are s t i l l not t o t a l l y "safe" characteristics, owing to the minute size of the plants. Spore size has been used i n the past as a distinguishing feature (Muller 1954, Schuster 197*0? but spore sizes for the two species i n B r i t i s h Columbia seemed to overlap. The most reliable distinguishing feature seems to be the nature of the elaters, i n particular, the width of the spirals. In A. julacea, elaters are 8-10 (im i n diameter, with 2(-3?) spirals which are 2 -4 |im wide. Elaters of A. juratzkana are 6-8 p.m i n diameter, with 2 -3 spirals that are only 2 |im wide. These differences i n the elaters were the primary 178 basis for identification of the two species of Anthelia i n this study, and a l l records shown on the maps of local distribution are based on species identified in this manner, at least for subalpine locations. ANTHELIA JULAGEA (L.) Dum. (See above key and generic description.) Sporophytes found on subalpine plants i n July through October. Distribution: Gircumboreal, Arctic-Alpine; i n the study area, Subalpine-Descending. Ecology: Growing in humid locations on i t s own humus, algal ooze, or s o i l over rock; on c l i f f faces of waterfalls or near the ocean. Substrate moist to wet with seepage, mist, or spray; usually open and northerly-facing. ANTHELIA JURATZKANA (Limpr.) Trev. (See above key and generic description.) Sporophytes found i n August. Distribution: Gircumboreal, Arctic-Alpine, Bipolar; i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: A pioneer on bare, exposed s o i l , compacted humus, or on s o i l over rock; occurring around permanent rock-pools i n areas of late snow-lie, on boulder-slopes, and on c l i f f s i n subalpine and alpine sites. Substrate mesic to moist, open. Family GEPHALOZIAGEAE1 The classification of genera by subfamilies given below follows Schuster (1972b, 1974). Schofieldia Godfr. (Godfrey 1976) i s placed i n Subfamily Gephalozioideae, i n which i t seems most closely related to 179 Pleuroclada (Hook.) Spruce. Subfamily HYGROBIELLACEAE HYGROBIELLA Spruce HYGROBIELLA LAXIFOLIA (Hook.) Spruce Plants minute to small, 240-880 \xm wide (the larger sizes on fertile female shoots), 5_20 mm high; dark to blackish-green, moss-like. Stems with flagella; cortical cells large, pellucid, forming a hyalodermis. Leaves transversely inserted, narrow, ovate-lanceolate; bilobed, with unequal lobes with the apices blunt to acute. Underleaves very large, similar in size and appearance to the leaves; bilobed or entire. Leaf cells elongated (to 70 |im long in leaf middle), with walls thin or evenly thickened, becoming golden to brownish; trigones lacking; cuticle smooth. Oil-bodies appearing absent, or, according to Schuster (1974), very small, several per cell, and finely segmented. Asexual reproduction lacking. Dioicous. Perianths slender, long-emergent, fusiform to cylindrical; the mouth denticulate with fingerlike cells. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, (oceanic-alpine); in the study area, Subalpine-Descending. Ecology: Essentially hygrophytic, growing on basic rock along cold streams in canyons, near waterfalls, and near the ocean at low elevations. Below the glacier on Mt. Matier, found on humus over an outcrop on an alpine slope, elevation ca. 1700 m. Subfamily GEPHALOZIOIDEAE 180 GEPHALOZIA (Dum.) Dum. Plants small, usually whitish to yellowish or clear green. Stems with distinct hyalodermis of much enlarged c e l l s ; postical branches and postical f l a g e l l a often present. Leaves succubously inserted, bilobed, the lobe apices acute. Underleaves lacking. Leaf c e l l s lacking trigones and oil-bodies; cuticle smooth. Gemmae occasionally produced. Inflorescences on postical branches. Perianth trigonous. Seta formed of 8 rows of outer c e l l s enclosing 4 rows of inner c e l l s . Sporophyte capsule wall 2-stratose. References: Underwood I896, Amakawa and Hattori 1952, Schuster 1964a. While the genus Gephalozia i s easily recognizable, Schuster (1974: 696-7OI) indicates that i t i s taxonomically d i f f i c u l t . Problems arise i n separation of species largely because of the small size of the plants, superficial similarities between species, and the microscopic nature of features essential to identification. Schuster also points out that knowledge of the sexual condition and nature of the perianth and bracts may be required at least for i n i t i a l identifications, u n t i l the facies of each species becomes familiar and st e r i l e plants can be relatively reliably identified. Schuster (1974: 70l) also states, "It i s always advisable to prepare stem cross sections i n identifying Cephalozia species, but i t must be kept i n mind that, with alterations i n vigor, considerable variation i n the number of c e l l rows i n both cortex and medulla may occur." For identification of species of Cephalozia i n this f l o r a a l l the features given i n the following descriptions were examined wherever possible, and the combination was then considered. Female bracts and perianths were often present on the plants. Sexual condition, while not 181 used as a primary criterion i n the following key, was determined wherever possible to corroborate evidence from other features of the species. 1. Leaves (when flattened) ovate, longer than wide; lobes straight or divergent; leaves not decurrent . . . . . . C. BICUSPIDATA 1. Leaves (when flattened) ± orbicular, as wide as long; lobes often incurved (connivent); leaves ± or longly decurrent 2 2. Cells at bases of leaf lobe small, usually less than 35 (am long; medullary c e l l s of stem usually i n fewer than 20 rows; dioicous; common on decaying wood C. LUNULIFOLIA 2. Cells at bases of leaf lobes larger, usually more than 35_40 |im long; medullary c e l l s of stem i n ca. 20 or more rows; autoicous; occurring i n bogs or i n subalpine meadows . . . . . . . 3 3. Leaves 8-12 cells broad, strongly decurrent; leaf lobes narrow, sharply pointed, 3 _ 5 c e l l s wide at base; cortical cells of stem i n 10-14 rows, medullary c e l l s i n 18-24 rows; perianth mouth c i l i a t e - l a c i n i a t e C. CONNIVENS 3. Leaves ca. 16-25 c e l l s broad, only slightly i f at a l l decurrent; leaf lobes broad, short-pointed, 5 _15 c e l l s wide at base; cortical c e l l s of stem i n 13-19 rows, medullary c e l l s i n 20-32 rows; perianth mouth crenulate to denticulate . . G. PLENICEPS GEPHALOZIA BICUSPIDATA (L.) Dum. Plants minute to small, 400-1000 p.m wide, pale whitish or yellowish-green or becoming golden- or red-brown. Branching postical and terminal-la t e r a l (branches then subtended by a 1-lobed l e a f ) . In transverse section, cor t i c a l cells of stem usually thick-walled, i n 8-16 (mean = l l ) rows; medullary cells usually thin-walled, occasionally also becoming thick-walled, varying i n plants from different locations from 6-24 rows (mean = 13). Leaves ovate, longer than wide, not decurrent dorsally, concave, the insertion subtransverse to oblique, 11-22 c e l l s wide. Leaf lobes narrowly triangular, 3-12 c e l l s wide at base; apices acute or acuminate. Cells at lobe bases thin- or thick-walled; small i n some plants, (12-) 16-26 x (16)24-36 p.m (mean = 20 x 28 (im); larger i n other 182 plants, (16-20)24-52 x 30-72 |im (mean = 30 x 44 p.m). Gemmae 1-celled, spherical; according to Schuster (1974) rare except i n subsp. ambigua. Autoicous. Female bracts bilobed with slender, lanceolate segments; margins entire (or subentire). Perianth mouth variable, crenulate to denticulate or dentate-setulose. Sporophytes very frequently found i n spring and summer. In this study, some simplification was necessary i n dealing with the genus Gephalozia, thus G. bicuspidata i s considered i n the broadest sense and not segregated into the various subspecies, e.g., ambigua, bicuspidata, and lammersiana. These subspecies represent differing states of ploidy (see Schuster 1974: 709), but are variously considered as separate species by other authors, and a l l three are l i s t e d as such by Schofield (1968b). The broad definition of G. bicuspidata followed here accounts for the great range of variation i n features. Distribution: Gircumboreal (the species s. l a t . ) ; i n the study area, Widespread (elevation-independent). Ecology: Occurring predominantly i n forests; also on ocean beaches, i n bogs, and i n subalpine meadows. Growing primarily on decaying wood (especially on logs, also on stumps and rotting wood on the ground); also frequently occurring on humus and sandy s o i l , on the ground or over rock. Substrate mesic, partially shaded to open. GEPHALOZIA CONNIVENS (Dicks.) Lindb. Plants minute to small, 400-1000 \m wide, pale golden-green. Branching postical. In transverse section of stem, cortical c e l l s thin-walled, i n 10-14 rows; medullary cells very thick-walled, i n 18-24 rows. Leaves orbicular, as wide as long, long-decurrent, ± f l a t , the insertion 183 nearly horizontal, 8-12 c e l l s wide. Leaf lobes long, narrowly triangular, connivent, 3 -5 c e l l s wide at base; apices sharply acute to acuminate. Cells at lobe bases thin-walled, 42-48 x 45-55 P-m« Gemmae ovate, apparently rare. [Above measurements taken from Schuster (1974); measurements from the one specimen collected i n the region of this f l o r a f e l l within these ranges.] Autoicous. Female bracts very deeply divided into 2-5 linear to lanceolate lobes. Perianth mouth c i l i a t e - l a c i n i a t e . Sporophytes not observed. Distribution: Circumboreal; i n the study area, Rare or Restricted, known only from the Delta Bog. Ecology: Growing on moist peat, shaded i n a hole, i n a raised, lodgepole pine - Sphagnum bog. CEPHALOZIA LUNULIFOLIA (Dum.) Dum. Plants minute to small, 450-800 p:m wide, pale whitish-green. Branching postical (ventral-intercalary). In transverse section of stem, corti c a l c e l l s thin-walled, i n 10-13 (mean = 11) rows; medullary cells thick-walled, varying among plants from different locations from 10-22(-26) rows (mean = 19). Leaves orbicular, as wide as long, 8-13(15) c e l l s wide, long-decurrent, f l a t , the insertion very oblique to nearly horizontal. Leaf lobes triangular, often connivent, 3-5(6) c e l l s wide at the base; apices short-acute. Cells at lobe bases thin-walled, 20-40 x 24-46(48) pja (mean = 28 x |im). Gemmae 1-celled, stellate. Dioicous. Female bracts bilobed, with entire margins. Perianth mouth crenulate-denticulate. Sporophytes found infrequently, May to July. Some variation was noted during identification of plants of this 184 species. Plants from the west coast of Vancouver Island seemed particularly luxuriant, with larger c e l l s and a rather distinctive facies resulting from more slender, more longly acute leaf lobes and a ± auriculate antical leaf base, as compared with plants from less humid sites away from the ocean. Distribution: Circumboreal; i n the study area, Widespread (essentially low-elevation). Ecology: Occurring predominantly i n forests, also on c l i f f s on ocean beaches. Growing primarily on decaying wood, especially on logs; occasionally found on humus on the ground; infrequently, on tree bases. Substrate usually mesic, partially shaded. CEPHALOZIA PLENICEPS (Aust.) Lindb. Plants small, 1000-1400 (im wide (the most robust species of Cephalozia occurring i n the study area); pure, deep green. Branching postical, and occasionally terminal-lateral. In transverse section of stem, cortical c e l l s thin-walled, i n 13-19(20) (mean = 16) rows; medullary c e l l s thick-walled, i n (13-17)20-32 rows (mean, without the low figures =27). Leaves broadly orbicular, at least as wide as long, (13~)16-32 cells wide, not or only slightly decurrent, ± concave, the insertion very oblique. Leaf lobes broadly triangular, not or l i t t l e decurrent, 5-15 c e l l s wide at base; apices short-acute or blunt. Cells at lobe bases thin-walled, (20)26-36(44, 48) x (24)30-56 |im (mean = 32 x 40 p.m). Gemmae ell i p s o i d a l , i n clusters at apices of attenuated shoots. Autoicous. Female bracts bilobed, with entire margins. Perianth mouth crenulate to denticulate. Sporophytes seemingly frequent, found from July to October. 185 Distribution: Gircumboreal - Missing i n Eastern Asia, Arctic-Alpine; i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Hygrophytic, subalpine, occurring i n meadows, on lake shores, i n swamps and bogs. Growing on humus or humusy s o i l ( t e r r e s t r i a l , or over rock), peat, and over Sphagnum; one specimen collected from a decaying log i n subalpine forest. Substrate moist to wet, usually open. GLADOPODIELLA Buch GLADOPODIELLA FLUITANS (Nees i n Funck) Joerg. Plants small to medium-sized, 1-2 mm wide, green with red pigments (plants from the study area), or becoming clear chestnut brown to blackish (according to the literature); creeping. Stems not differentiated, cortex not forming a hyalodermis; cortical c e l l s differing l i t t l e i n diameter from medullary c e l l s . A l l branches, including leafy and stoloniferous, postical-intercalary i n origin. Rhizoids sparse, scattered. Leaves succubously (to subhorizontally) inserted, distant to almost contiguous, oblong-elliptical, widest above middle; somewhat asymmetrically bilobed, the sinus descending 0.2-0.4 x the length of the leaf; lobe apices obtuse to broadly rounded. Marginal c e l l s of leaf ca. 30-35 pJn long; median c e l l s usually ca. 30-40 x 35-^5 pjri; basal cells to 38-42 x 38-60 (im [measurements from Schuster (l9?4: 6 8 l ) ] . Cell walls thin or slightly thickened; trigones lacking; cuticle smooth. Oil-bodies 4-8 per c e l l , finely granular, ca. 4.0-5-5 "to 4.0-5.5 x 6-10 [im (Schuster 1974: 681). Underleaves distinct under compound microscope, 3~8 c e l l s broad at base, variable, lanceolate, subulate, or bilobed, with stalked marginal slime papillae. Asexual reproduction absent. Plants from the study area were s t e r i l e . The species i s dioicous. Gynoecia on short postical branches. 186 Distribution: Circumboreal; i n the study area, Rare or Restricted (known from Lulu Island, Richmond, just south of Vancouver; WBS 26208). Ecology: Growing over Sphagnum i n a bog ditch adjacent to a railroad track. The ditch preserves a remnant of a bog flo r a formerly more extensive i n the area. PLEUROCLADA Spruce PLEUROCLADA ALBESCENS (Hook.) Spruce Plants small, 640-880 |im wide; clear green without secondary pigmentation; ± julaceous. Stems lacking fl a g e l l a ; i n transverse section, cortical c e l l s very large, thin-walled, forming a hyalodermis. Leaves strongly concave, bilobed, round-quadrate; the antical half transversely inserted. Underleaves large', subequal to leaves i n length; ovate-lanceolate, occasionally with basal tooth. Cells lacking trigones and oil-bodies; walls thin, or equally thickened; cuticle smooth. Asexual reproduction rare, by green, 1-celled, spherical to e l l i p t i c a l gemmae (Schuster 19?4, Schuster and Damsholt 19?4, Paton 19?6); not observed i n plants from the area of this f l o r a . Dioicous. Plants from the study area were s t e r i l e . Distribution: Circumboreal, Arctic-Alpine; i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing i n open subalpine meadows on s o i l and humus, over the ground or over rock outcrops, c l i f f faces, slopes, or stream banks. Substrate moist, becoming wet with flowing water and seepage from melting snow. References: Szweykowski and Krzakowa 1966, Krzakowa I969. 187 SCHOFIELDIA Godfr. SCHOFIELDIA MONTICOLA Godfr. Plants large, 1.5~3«0 mm wide, 8-15 mm long; clear, bright green, lacking secondary pigmentation; succulent, growing i n dense turfs. Stems whitish-green, fleshy, and b r i t t l e ; i n transverse section, the thin-walled cort i c a l c e l l s similar to the medullary c e l l s i n appearance; hyalodermis lacking. Leaves succubously inserted, becoming transverse dorsally; somewhat broader than long, r i g i d , concave; bilobed, occasionally trilobed or with a hook on the antical base. Underleaves frequent, though usually obscured by rhizoids, which are scattered along the ventral stem; variable i n size and shape, not exceeding l/3 the size of the leaves. Leaf c e l l s thin-walled; trigones essentially lacking; cuticle smooth. Median leaf c e l l s 30-52 x 40-94(100) (im; marginal c e l l s 20-48 x 30-62 (jjn. Oil-bodies absent. Asexual reproduction by (l-)2-celled, green, e l l i p s o i d or ovoid gemmae, 14-20 x 20-34 [im; produced among the youngest leaves at the stem apex. Dioicous. Gynoecia terminal on leafy stems or borne on short l a t e r a l (?) branches. Bracteole present. Perianth plicate; mouth lobulate with 1-2-celled teeth. Sporophytes found infrequently, August through October. Seta 4 c e l l s i n diameter, consisting of 8 rows of epidermal c e l l s and 4 rows of inner c e l l s . Capsule wall 2(-3)-stratose. For the complete description of this species and genus, with i l l u s t r a t i o n s and discussion, see Godfrey (1976). Distribution: Endemic to Western North America; i n the study area, S t r i c t l y Subalpine-Alpine. Ecology: Growing on seepy slopes or banks of snow-melt r i l l s , 188 streamlets, or ponds, sometimes ± exposed, but usually partially to completely shaded by heathers, i n subalpine meadows at elevations of approximately 1200-2000 m. Subfamily 0D0NT0SCHISMAT0IDEAE 0D0NT0SCHTSMA (Dum.) Dum. Plants small to medium-sized, 0.8-1.5 mm wide, up to 3 or more cm long; yellow-green, becoming reddish or brownish. Stems prostrate, the shoot apices ascending. Branching mostly postical (occasionally l a t e r a l ) ; leafless, postical f l a g e l l a frequent on stems. Rhizoids scattered. Leaves succubous, orbicular-ovate, entire. Underleaves present, though often minute and d i f f i c u l t to see; usually flap l i k e , with slime papillae on margins. Leaf c e l l s with very coarsely bulging, often confluent, trigones, the c e l l lumen becoming stellate. Oil-bodies usually 2-4 per c e l l , granular, large, almost f i l l i n g c e l l lumen. Usually dioicous (Schuster 1974). Plants from the study area s t e r i l e . References: Evans 1903-Jamesoniella autumnal!s and Jungermannia spp. may be mistaken for plants of this genus, especially i n sterile specimens. 1. Leaf c e l l s with middle lamella indistinguishable, no more pigmented than the c e l l walls; cuticle ± papillose; leaves rather strongly concave; plants bright reddish-brown; gemmae frequent, on 3 _ranked flagelliferous shoot apices . . . 0. DENUDATUM 1. Leaf c e l l s with middle lamella obvious, more distinctly pigmented (golden to brownish) than the c e l l walls and trigones; cuticle smooth; leaves only barely concave; plants golden-brown, becoming fuscous; gemmae infrequent 0. ELONGATUM 189 ODONTOSGHISMA DENUDATUM (Nees i n Mart.) Dum. Plants 1-2 mm wide, light yellow-green to golden- or red-brown. Shoots frequently ± abruptly terminated by flagelliferous apices bearing reduced or no leaves. Leaves somewhat to strongly concave. Underleaves usually minute; occasionally becoming large and distinct near base of a branch; equal i n size to the leaves on gemmiparous shoot apices. Middle lamella obscure. Gemmae frequent, 2-celled, green to pinkish-red or purple, variable i n size, e l l i p t i c a l and 12-16 x 18-28 |im, or spherical and smaller; produced on margins of transversely inserted leaves and underleaves (which then become erose-dentate) on 3_ranked, virtu a l l y radially symmetrical flagelliferous shoot apices. There i s apparently much variation i n Odontoschisma denudatum, and this, I believe was the basis for Evans' (1903) description of a new species, 0 . gibbsiae from Port Renfrew, Vancouver Island. Evans (1903) separates 0. gibbsiae from 0. denudatum largely on the basis of the suberect, strongly concave, and imbricated leaves and underleaves of the gemmiparous branches (squarrose, plane or slightly concave, and distant i n 0, denudatum), and on thick-walled gemmae (supposedly thin-walled i n 0. denudatum). He then separates 0. gibbsiae from 0. macounii on the basis of i t s frequent reddish to brownish coloration and smaller l e a f - c e l l size. Evans' il l u s t r a t i o n s of 0. gibbsiae show large, distinct underleaves on a st e r i l e , nongemmiparous stem; such large underleaves are, according to his description, found at the base of a branch. On the basis of study of several specimens of Odontoschisma from the west coast of Vancouver Island, i t seems that 0. gibbsiae represents one phase of variation i n a polymorphic species, i.e., 0. denudatum. In 0. denudatum s. l a t . , then, are included plants with compacted, gemmiparous apices with concave, imbricated leaves and underleaves, and also, slender, attenuate gemmiparous apices, with distant, squarrose or plane leaves and underleaves. Transitions between, and combinations of, both forms can be observed i n the same mat of Odontoschisma collected from a single site. I t seems that production of flagelliferous stem apices and attenuated, distant-leaves gemmiparous apices may be environmentally induced by such factors as greater exposure to light (and, thus, heat) and/or less available moisture, as, for example, i n plants growing on rotting stumps i n open bogs where the substrate i s exposed and subject to drying. The compact, imbricate-leaved gemmiparous shoots may be produced under more "favorable" conditions involving greater shade and moisture, e.g., i n the rainforest. Underleaves on the plants studied were found to vary i n size. The underleaves are typically minute and most easily observed near the stem apices, but i n a few specimens, large underleaves l i k e those i l l u s t r a t e d by Evans (1903s Figure 30) for 0. gibbsiae, were observed near the bases of branches. In one specimen (JDG 27_7_6) the tuft of plants i n which this feature was noted also contained shoots with naked, flagelliferous shoot apices, a feature frequent i n 0. denudatum. Distribution: Disjunct Western North America — Eastern North America — Western Eurasia — Eastern Asia, Bipolar (?); i n the study area, S t r i c t l y Oceanic. Ecology: Growing on decaying logs and stumps in the forest; on peaty humus on tree bases and over decaying stumps i n bogs; and on humus on the ground on a rocky promontory by Kennedy Lake, Substrate mesic to moist, occasionally becoming wet; p a r t i a l l y shaded to open. 191 ODONTOSCHISMA ELONGATUM (Lindb.) Evans Plants ca. 1.0-1.5 nun wide; green to golden-brown, with a tendency toward fuscous coloration. Shoot apices never (Schuster 197*0 becoming flagelliferous. Leaves laxly imbricate, l i t t l e concave. Underleaves small to distinct along entire postical stem surface. Middle lamella conspicuous, becoming golden or brownish, darker and more dist i n c t l y pigmented than the c e l l walls. Gemmae not observed; gemmiparous shoots, according to the literature, short, with little-reduced leaves. Distribution: Disjunct North America — Western Eurasia; i n the study area, Hyperoceanic. Ecology: Growing on i t s own humus over sand and algal ooze on NW-facing c l i f f faces, above a beach on the Pacific Ocean (Brooks Peninsula). Substrate moist, exposed. Family CEPHALOZIELLACEAE CEPHALOZIELLA (Spruce) Schiffn. i n Engler et Prantl Plants minute, 1^5-300 \xm wide ( s t e r i l e portions of stems), 1-8 mm long, threadlike. Cortical c e l l s of stem ca. the same size as the inner c e l l s . Rhizoids few, scattered along the stem. Branching late r a l or postical. Leaves usually distant, transversely inserted, bilobed for 0.50-0.75 x their length. Leaf lobes varying among the different species from 2 to 9 cel l s wide at the lobe base. Oil-bodies present i n leaf cells, roughly spherical, 2-3 |im i n diameter. Underleaves present or absent. Gemmae frequently present, at stem apices or on uppermost leaves (and underleaves). Plants dioicous, autoicous, or paroicous. Perianths long-emergent, 192 ± plicate, terminal on short or long postical shoots. Bracteole present. Sporophyte with capsule wall bistratose, seta consisting of 4 c e l l rows i n transverse section. References: Arnell 1950; Kitagawa 1962a, 1965a; Schuster 1971b. Schuster (1953s 482) states, "The members of this genus are the most d i f f i c u l t of Hepaticae to identify, partly because of their minute size, partly because of the chaotic condition of their taxonomy." Schuster and other authors (Macvicar 1926, Muller 1956, Schuster and Damsholt 1974) use sexual condition as an important criterion i n species identification. In Gephaloziella, most species are either dioicous or autoicous and determination of either condition i s made very d i f f i c u l t by the minute and delicate nature of the individual shoots. A number of shoots must be carefully separated from the mat of plants; individual shoots must be further separated without tearing off sexual branches. The key below i s not based primarily on sexual condition, but uses more easily discernible features of the plants. To ensure accuracy i n species identification, however, one should attempt to determine the sexual condition wherever possible. 1. Margins of nongemmiparous leaves entire 2 1. Margins of nongemmiparous leaves dentate 4 2. Underleaves lacking on st e r i l e , nongemmiparous shoots; bracteoles lacking on f e r t i l e female shoots . . . . (See Eremonotus myriocarpus or Marsupella) 2. Underleaves on s t e r i l e , nongemmiparous shoots easily distinguished; bracteoles present on female shoots 3 193 3. Leaf margins entire; abaxial surfaces of leaves smooth, lacking cuticular protuberances C. DIVARICATA 3. Leaf margins entire to crenulate or somewhat dentate, abaxial surfaces of leaves with cuticular bumps or protrusions . . . . C. DIVARICATA var. SCABRA 4. Margins of leaves with a few isolated teeth 5 4. Margins of leaves strongly dentate, with many teeth or spines . . . 6 5. Leaf lobes 6-10 c e l l s wide at base; leaf margins ± entire, crenulate or with a few small teeth; abaxial leaf surface with rather large cuticular bumps