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The bryophyte flora of Bridal Veil Falls, British Columbia : an analysis of its composition and diversity Djan-Chekar, Nathalie 1993

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THE BRYOPHYTE FLORA OF BRIDAL VEIL FALLS, BRITISH COLUMBIA:AN ANALYSIS OF ITS COMPOSITION AND DIVERSITYbyNATHALIE DJAN-CHEKARB.Sc., McGill University, 1990A THESIS SUBMITTED IN PARTIAL FULFILLMENT OFTHE REQUIREMENTS FOR THE DEGREE OFMASTER OF SCIENCEinTHE FACULTY OF GRADUATE STUDIES(Department of Botany)We accept this thesis as conformingto the required standardTHE UNIVERSITY OF BRITISH COLUMBIAApril 1993Copyright, Nathalie Djan-Chekar, 1993In presenting this thesis in partial fulfilment of the requirements for an advanceddegree at the University of British Columbia, I agree that the Library shall make itfreely available for reference and study. I further agree that permission for extensivecopying of this thesis for scholarly purposes may be granted by the head of mydepartment or by his or her representatives. It is understood that copying orpublication of this thesis for financial gain shall not be allowed without my writtenpermission.Department of BotanyThe University of British ColumbiaVancouver, CanadaDate^5 A-v-11i 111'3(Signature) DE-6 (2/88)AbstractBridal Veil Falls is a small provincial park located at the eastern extremity of theFraser Lowland in southwestern British Columbia. This study presents an inventory of thebryophyte flora of Bridal Veil Falls Provincial Park and an adjacent boulder slope. Thisinventory provided the basis for a discussion of the ecology and geographical affinities of thebryoflora emphasizing the environmental and historical factors that are determinant of itscomposition. A key to the mosses is provided.Field work conducted between 1991 and 1992 and examination of herbariumspecimens revealed a diverse bryoflora in comparison to areas of similar size in BritishColumbia's Lower Mainland. This bryoflora is composed of 210 species (150 mosses and 60liverworts). In addition, Barbula convoluta var. gallinula was found new to British Columbia.An analysis of bryophyte occurrence in habitats showed that high species diversity canbe attributed to the wide variety of both habitats available at the site and microenvironmentsassociated with these different habitats. Variations of light, moisture, and edaphic conditionswere found to be generally important sources of diversity.The bryoflora of Bridal Veil Falls has a strong circumboreal component. Its affinitywith the rich bryoflora of coastal British Columbia is reflected by a large proportion ofPacific North American endemics and species of humid coastal regions. The availability ofsuitable microenvironments allows the occurrence of species of arctic-montane, meditenaneanand strictly oceanic distribution. The bulk of the flora is probably derived from bryophytepopulations that colonized the Lower Mainland of British Columbia from southern refugiaafter the retreat of the Wisconsin ice.IITable of contentsAbstract ^  iiList of tables  viList of figures ^  viiAcknowledgement  ixIntroduction  ^11.1. Objectives  ^11.2. Description of the study area ^  31.2.1. Physiography, geology and geomorphology. ^ 31.2.2. Climate  ^81.2.3. Vegetation  ^91.3. Late-Pleistocene and recent history ^  121.3.1. Geology ^  121.3.2. Vegetation and climate ^  131.4. Earlier bryological work at Bridal Veil Falls ^  16Inventory and keys ^  172.1. Materials and methods ^  172.1.1. Collection  172.1.2. Identification and preparation of the keys ^  172.1.3. Nomenclature ^  191112.2. Results and discussion ^  202.2.1. Diversity of the flora  202.2.2. Keys to the mosses of Bridal Veil Falls Provincial Park andadjacent boulder slope. ^  212.2.3. Notes on problematic taxa  492.2.4. Barbula convoluta var. gallinula, new to British Columbia ^ 54Ecology ^  553.1. Introduction - Species diversity ^  553.2. Materials and methods  563.2.1. Definition of habitat categories ^  563.2.2. Definition of substratum categories  573.2.3. Similarity indices ^  583.3. Results and discussion  593.3.1. Floristic diversity in terms of habitats and substrata ^ 593.3.2. Widely distributed versus restricted species ^ 613.3.3. Floristic similarity between habitats  663.3.4. The occurrence of species in habitats and on substrata ^ 703.4. Conclusion ^  81Phytogeography  824.1. Materials and methods ^  824.1.1. Defmition of phytogeographic elements ^  82iv4.2. Results and discussion ^  854.2.1. Affinities of the flora - broad geographic perspective ^ 854.2.2. Affinities of the flora - regional geographic perspective. ^ 994.2.3. Relationship between habitat and geographic affinity. ^ 1024.2.4. An interpretation of the development of the flora.  109Literature cited ^  112Appendix A. Annotated list of mosses at Bridal Veil Falls ^  121Appendix B. Annotated list of liverworts at Bridal Veil Falls  125Appendix C. Occurrence in habitats and on substrata of species collected at BridalVeil Falls during the course of this study. ^  127Appendix D. Synonyms and alternative references.  131Appendix E. Similarity between habitat/substratum combinations based on composition oftheir respective moss flora using the Jaccard Index of Similarity. ^ 133Appendix F. A Note of caution^  134List of TablesPagesTable 3.1.^The distribution of bryophytes between habitats, between substrata, andbetween substrata within each habitat. ^  60Table 3.2.^Number of bryophytes unique to each habitat. ^  63Table 3.3.^Number of bryophytes unique to each substratum  64Table 3.4.^Similarity between habitats based on composition of the moss flora using theJaccard Index of Similarity. ^  67Table 4.1.^Phytogeographic elements based on distributions within the NorthernHemisphere. ^  83Table 4.2.^Phytogeographic elements based on regional (Pacific North American)distributions. ^  84Table 4.3.^The number of bryophytes belonging to each circumboreal phytogeographicelements ^  86Table 4.4.^The number of bryophytes belonging to each regional phytogeographicelements ^  100Table 4.5.^Number of bryophytes species of each regional phytogeographic elementoccurring in each habitat type. ^  103viList of FiguresPagesFigure 1.1.^Location of Bridal Veil Falls Provincial Park, British Columbia ^ 2Figure 1.2.^Schematic diagram of the study area ^  4Figure 1.3.^Physiographical setting of Bridal Veil Falls. ^  5Figure 1.4.^Bedrock geology of the study area, 1:25 000.  7Figure 1.5.^Temperature and precipitation at Agassiz, British Columbia. ^ 10Figure 1.6.^Schematic diagram of the pattern of land, sea, and ice during the Wisconsinglacial maximum ^  14Figure 3.1.^Relationship between habitats based on similarity in species composition usingmultidimensional scaling. ^  68Figure 4.1. World distribution of Hylocomium splendens. ^  87Figure 4.2. World distribution of Hymenostylium insigne  90Figure 4.3.^World distribution of Antitrichia californica ^  91Figure 4.4.^Claopodium crispifolium, a) world distribution, b) Pacific North Americandistribution. ^  93Figure 4.5. Hypnum subimponens, a) world distribution, b) Pacific North Americandistribution ^  94Figure 4.6.^Pseudotaxiphyllum elegans, a) world distribution, and Stormer 1969), b) PacificNorth American distribution. ^  95viiFigure 4.7. Pacific North American distribution of the endemic Racomitrium lawtonae.^  97Figure 4.8. Pacific North American distribution of the endemic Pseudobraunia californica.^  98Figure 4.9. Regional distribution of a) Chandonanthus fihformis, b) Moerckia hibernica,and c) Odontochisma denudatum ^  106viiiAcknowledgementI would like to thank my research supervisor, Dr. Wilf Schofield, for hisencouragement and assistance throughout this work. I am truly grateful for the kind mannerwith which he has shared his tremendous wealth of bryological knowledge with me.I would also like to thank my committee members, Drs. Gary Bradfield and GilbertHughes, for their help and support.Rend Belland's sharing of knowledge and assistance with ecological analyses wasgreatly appreciated. Much thanks also to Olivia Lee for herbarium assistance. I am alsograteful to Alan Reid for his companionship and help in field collection.Expert opinion for the identification of Barbula convoluta var. gallinula andScleropodium touretii var. colpophyllum was graciously provided by Dr. R.H. Zander and Dr.Enroth, respectively. Dr. J.W.H. Monger and Dr. K.W. Savigny provided useful informationabout the geology of the study area.Poet Anton Baer proof-read the manuscript.This research was partially funded by a scholarship from the Natural Sciences andEngineering Research Council of Canada.Finally, I would like to thank the British Columbia Ministry of Parks for theopportunity to work at Bridal Veil Falls.ixChapter 1. Introduction1.1. ObjectivesFloristic studies provide the foundation for an understanding of biodiversity in plants.They also present the background for studies of the environment. In British Columbia'sProvincial Parks, such detailed studies are rare, and there are none of bryophyte diversity.Bridal Veil Falls Provincial Park was chosen for its ready accessibility, and because earlierbryophyte collections made there suggested an exceptional diversity for its size and presenteda number of intriguing phytogeographical problems. The aim of this study was thus to preparea detailed documentation of the bryophyte flora of Bridal Veil Falls area with consideration ofthe environmental and historical factors that determine its composition.Bridal Veil Falls Provincial Park is located in the coastal region of British Columbiaabout 100 km east of Vancouver in the Lower Fraser River Valley (Fig. 1.1). The climate ofcoastal British Columbia is especially favourable for bryophytes (Schofield 1992). It ischaracterized by year-round moderate temperatures, high rainfall and mild winters (Hare andThomas 1974). Although the area was almost completely covered by ice during the Wisconsinglaciation, favourable coastal refugia south of the glacial boundary and along the southerncoast of Alaska helped preserve this rich bryoflora (Schofield 1984, 1992). The mild andhumid coastal climate and historical circumstances combine with the variety of microhabitatsavailable at Bridal Veil Falls to create a high potential for an abundant and rich bryophyteflora.1col>KAMLOOPS•n0 VANCOUVER HOPEBRIDAL VEIL FALLSCANADAUSAFigure 1.1. Location of Bridal Veil Falls Provincial Park, British Columbia.2Specific objectives of this study were:1. To present an inventory of the bryophytes of Bridal Veil Falls area.2. To design a key to the mosses of the area.3. To describe and discuss bryophyte occurrence in the various habitats andsubstrata found at the site.4. To analyse geographical affinities of the species composing the flora.5. To examine the relationship between habitat and geographic affinity.6. To examine bryophyte diversity at Bridal Veil Falls in light of late-Pleistoceneand Holocene history of coastal British Columbia and the Fraser Lowland.1.2. Description of the study areaBridal Veil Falls Provincial Park was established in 1965 and comprises an area of 32hectares (British Columbia Parks Branch 1990). The study area includes the park, and aboulder slope immediately adjacent to the west of the park. The boulder slope was included inthe study area because it represents a habitat very different from those found in the park. Italso appears of particular bryological and phytogeographical interest based on earliercollections. The whole of the study area therefore covers approximately 60 hectares. Figure1.2 presents a schematic diagram showing the different environments found in the study area.1.2.1. Physiography, geology and geomorphology.Bridal Veil Falls is located near the eastern extremity of the Fraser Lowland in BritishColumbia's Lower Mainland (Fig 1.3). The Fraser Lowland belongs to the Georgia3Nroad,--, -----cv ') picnic areanr-1Figure 1.2. Schematic diagram of the study area (not to scale).4Figure 1.3. Physiographical setting of Bridal Veil Falls. (from Campbell et al. 1990)5Depression, a large basin between the highlands of Vancouver Island and the coastalmountain ranges of the mainland (Campbell et al. 1990). The Fraser Lowland is a low lyingarea of depositional origin between the Coast mountain range to the north and the Cascademountain range to the south (Holland 1976). Bridal Veil Falls is at the base of the lowland'ssouthern escarpment. The study area is thus bordered to the south by northwest facing cliffs(Fig. 1.2). It lies between 100 and 400 meters above sea level.The bedrock geology of the area of Bridal Veil Falls is very complex. An outline ofthe nature of the bedrock in the study area and immediate surroundings is presented in Figure1.4. The cliffs found in the study area are predominantly of sedimentary calcareous rock.They are outcrops of the zone of limestone seen in Figure 1.4. Non-calcareous cliffs arefound above the boulder slope. Rocks in the boulder slope are also non-calcareous.The boulder slope probably represents one of the rockfalls that Armstrong (1984)described as common along the Trans-Canada highway east of Rosedale. These rockfallsconsist of free fall or precipitous movement of fragments of bedrock of various sizes detachedfrom cliffs or very steep slopes. Rockfalls are usually caused by freeze-thaw action. It ispossible that the process resulting in the boulder slope at Bridal Veil Falls was not sudden.Rock fragments detached from the cliff above could have accumulated over a period of time,and this could still be happening (K.W. Savigny, pers. communication). It is clear, however,that the bulk of the rockfall near Bridal Veil Falls had already occurred by 1963; the boulderslope is visible on air photos taken that year.Four waterfalls are found along the cliffs in the study area (Fig. 1.2). In several places,as well, water dribbles over the rock faces. Four streams are associated with the waterfalls, at614`)Figure 1.4. Bedrock geology of the study area, 1:25 000. (modified from Monger 1966)v‘"--^, ^-, 13RI2AL VEIL FALLS--'r PRovirslcIAL PARK^,, ^t^, 1 I,^. 1,^1, I,^i If;/ I^1k^1^/'k :^/^I■.1.4.'" .......i ‘'./.^ <0green,is liceous^..." / / "f<^VolcaYlic. rock ------/--- / 7^skate.^cs.1 1 & 1Sabia.stone,iime&forie.Jelineation of -file .stuol y area/./^pprOXi mofe, 10061-1011 Of the h)ovilcier Slope,7least one of which is intermittent. The water is likely to have a very high content of calciumcarbonate. There is evidence for this in the abundant marl deposits found along the FraserRiver below Bridal Veil Falls (Smith 1971; J.W.H. Monger, Geological Survey of Canada,pers. communication). In the form of calcite, calcium carbonate is a main constituent oflimestone (Cole 1983). Calcium could be dissolved into the water as it flows over thelimestone outcrops (Fig. 1.4). Furthermore, above Bridal Veil Falls on the slope of CheamPeak are other areas of limestone and large areas of shales and sandstones of the CultusFormation (not shown in Figure 1.4). These shales and sandstones contain calcium feldspar(J.W.H. Monger, pers. communication). In addition, the large amount of rainfall in the areaand the cool temperatures of the northwest-facing mountain slope are conducive to thedissolution of relatively important quantities of calcium into the water (K.W. Savigny, pers.communication). Solubility of calcium carbonate increases as temperature falls from 35°C to0°C (Cole 1983).1.2.2. ClimateBridal Veil Falls is located in the Coastal Western Hemlock zone as defined by thebiogeoclimatic system of ecosystem classification of British Columbia. This zone lies at lowto middle elevations all along the coast of British Columbia, and penetrates the coastalmountain barrier in the Fraser River Valley (Pojar et al. 1991). It is characterized as therainiest zone in the province. Typically, it has cool summers and mild winters, with meanannual temperatures of about 8°C (Pojar et al. 1991). The growing season in the Lower FraserRiver Valley is among the longest in British Columbia (Campbell et al. 1990).8Figure 1.5 presents data from a climatic station in Agassiz, on the opposite side of theFraser River from Bridal Veil Falls. The mean monthly temperatures peak at 18°C duringJuly and do not fall below 0°C. Snowfall is light, with snow accumulation reaching a meanmonthly maximum of around 35 cm in January. Precipitation is highest during the fall andwinter, and lowest during the summer. Nevertheless, more than 80% of the plants' waterrequirements are met by precipitation during the growing season (Farley 1979). Within thestudy area itself, the northwest-facing escarpment undoubtedly exaggerates the cool summertemperatures, especially near the cliff bases, by reducing periods of direct sunlight. The cliffsthus remain cool and mesic throughout the year.1.2.3. VegetationThe forest of the study area has been logged during the past century. Most of the treesare estimated to be between 35 and 50 years old (S. Doubinin, British Columbia Ministry ofForest, pers. communication). The area would thus have been logged in the 1940's and1950's, not long before the creation of the park in 1965. The British Columbia Ministry ofForest has no record of major fires in the area (S. Doubinin, pers. communication). Burntstumps scattered in the forest are likely to be the remains of individual trees struck bylightning.Two tree species dominate the forest: bigleaf maple (Acer macrophyllum Pursh) andwestern redcedar (Thuja plicata Donn ex D.Don). They serve to support an abundantepiphytic flora of bryophytes, lichens and licorice fern (Polypodium glycyrrhiza D.C. Eat.).9Figure 1.5. Temperature and precipitation at Agassiz, British Columbia. (data from Farley1979)20 - 15 -10 -5 -o —J F M AM J J AS ON D=MD ON. ORD mean monthly temperaturemean monthly precipitation_CL mean snow accumulation-35-30-25- 20- 15-10- 510Other tree species found in the area include western hemlock (Tsuga heterophylla (Raf.)Sarg.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), paper birch (Betula papyriferaMarsh.), red alder (Alnus rubra Bong.), and black cottonwood (Populus trichocarpa Torr. &Gray). The understory is rather sparse. Vine maple (Acer circinatum Pursh) is common, as areferns, notably the western sword fern (Polystichum munitum (KauIf.) Presl.), and near thecliffs and streams, the maidenhair fern (Adiantum pedatum L.). Under clearings in the canopy,shrubs such as devil's club (Oplopanax horridum (J.E.Sm.) Miq.), salmonberry (Rubusspectabilis Pursh) and thimbleberry (Rubus parviflorus Nutt.) are abundant.The dominant vegetation in the boulder slope is of bryophytes. In the open portions ofthe slope, Selaginella wallacei Hieron. is common. Other vascular cryptogatns to be foundthere are mainly: Huperzia myoshiana (Makino) Ching, Asplenium trichomanes L.,Cryptogramma acrotichoides R.Br. in Richards, Woodsia R.Br. spp., and Polypodiumglycyrrhiza which flourishes during the rainy season. Among flowering plants, Geraniumrobertianum L., and three species of miner's lettuce (Montia parvtfolia (Moc. ex DC.)Greene, Claytonia perfoliata Donn ex Willd., and C. sibirica L.) are common. Ocean spray(Holodiscus discolor (Pursh) Maxim.), false azalea (Menziesii ferruginea Smith), Douglasmaple (Acer glabrum Ton.), stinging nettle (Urtica dioica L.), and Ribes L. spp. dominate theshrubby areas.111.3. Late-Pleistocene and recent history1.3.1. GeologyDuring the last glaciation, the Cordilleran ice sheet occupied most of BritishColumbia. In southwestern British Columbia, the ice flowed eastward from the VancouverIsland ranges and westward from the Coast mountains (Holland 1976). The Fraser Lowlandwas entirely covered by ice by around 17 000 B.P., as the ice sheet reached the Vancouverarea (Fulton 1984). At glacial maximum, the ice extended to the Puget Lowland, about 250km south of British Columbia's Lower Mainland (Mullineaux et al. 1965, as reported inFulton 1984). Thirteen thousand years B.P., the retreating ice allowed the sea to invade theStrait of Georgia (Clague 1981; Fulton 1984). While the sea was invading the western part ofthe Fraser Lowland, in the eastern part, there was a short, local re-advance of a piedmontglacier referred to as the Sumas advance (Clague 1981). Piedmont glaciers are the spreading,terminal part of valley glaciers that occupy broad lowlands at the base of steep mountainslopes (Flint 1971). The Sumas glacier eroded the mountain slopes in which it was embedded.The steep cliffs found along the base of the escarpments bordering the Fraser Lowland, suchas those found at Bridal Veil Falls, are the products of erosion by this glacier (K.W. Savigny,pers. communication). The Sumas glacier receded rapidly from the Fraser Lowland. Thelowland was then probably free of glacial ice by 11 000 years B.P. (Clague 1981). Followingthe retreat of the ice, the eastern portion of the lowland was not invaded by the sea; the driftdeposited by the Sumas glacier was not overlain by marine sediments (Clague 1981).More recently, the mountain slope at the base of which Bridal Veil Falls is locatedwas the site of a major rock avalanche. This avalanche, referred to as the Cheam Slide,12occurred at some time between 4 300 and 5 100 years B.P. (Naumann and Savigny 1991). Itsdebris overlies the Fraser River floodplain (Smith 1971). The source was located just aboveBridal Veil Falls at an elevation of 1 200 m on Cheam Peak (Smith 1971, as reported inArmstrong 1984).1.3.2. Vegetation and climateDuring full-glacial times, two major ice-free areas could have served as refugia for theplants that colonized the glaciated lands after the retreat of the ice (Fig. 1.6). To the south ofthe ice, most of the United States were ice-free. To the north, ice-free conditions existed overmajor portions of Alaska and Yukon, which were united to the Siberian peninsula by theBering Land Bridge (Ritchie 1987). On a smaller scale, numerous lowland refugia arebelieved to have been scattered along the shores and the offshore islands of Alaska andBritish Columbia (Pielou 1991). For example, geological and biological evidences have shownthat small areas in the Queen Charlotte Islands probably served as refugia during the lastglaciation (Heusser 1960; Calder and Taylor 1968; Schofield 1969a, 1989; Mathewes et al.1982, as reported in Hebda 1983; Pielou 1991). Some bryophytes could also have survivedthe glaciation in high mountain refugia, or nunataks (Pielou 1991).After the retreat of the ice, it is probable that lowland coastal vegetation promptlyinvaded southwestern British Columbia. Considering the region's proximity to thesouthernmost extent of the ice, the most likely source of propagules was southern refugia.Plants of the modern west coast flora could have survived glaciation quite close to the ice13Figure 1.6. Schematic diagram of the pattern of land, sea, and ice during the Wisconsinglacial maximum. The fine lines show modern coastline. (from Pielou 1991)Ice-free^Frozen sea^Ice sheetland (frozen mostof the year)14margin, possibly in valleys protected from the generally harsh climatic conditions (Barnosky1985, as reported in Pielou 1991). In mountainous regions, very different climates can prevailin adjacent areas. As well, the proximity of the ocean was likely to attenuate climaticextremes.Around 10 500 years B.P., temperature rose rapidly and precipitation declined, suchthat between approximately 10 000 B.P. and 7 500 B.P., conditions in the Lower Mainlandwould have been generally warmer and drier than at present (Mathewes 1973; Mathewes andHeusser 1981). The effect of this so-called, xerothermic period (the Hypsithermal) onvegetation is not clear, however. According to Mathewes (1973), the effect of macroclimaticchanges on vegetation during the Hypsitherrnal could have been attenuated by the widetolerance ranges of most tree species and the ameliorating oceanic influence along the coast.For example, in Hypsithermal deposits from the Lower Mainland, Mathewes (1973) foundmacrofossil of species of mosses characteristic of humid regions of the coast and interior. Hisfindings support the arguments for the prevalence of humid coastal conditions in the FraserLowland since the retreat of the ice. It is also possible that humid conditions existedthroughout the Hypsithermal as a microclimate in protected sites.From 7 500 to 6 000 B.P., a trend toward cooler and wetter microclimatic conditionsmarked the end of the xerothermic period. Since then, climatic conditions in the region appearto have been more or less constant (Mathewes and Heusser 1981).151.4. Earlier bryological work at Bridal Veil FallsThroughout the years, Bridal Veil Falls has been extensively collected for bryophytes.Most of the specimens are housed at the herbarium of the University of British Columbia(UBC). As early as the 1940's, V.J. Krajina collected in the area and it is possible he visitedthe site later to become Bridal Veil Falls Provincial Park. Early specimens from the BridalVeil Falls proper date from the 1960's. The collections are mainly those of W.B. Schofield,who first collected at the site in 1964. Among his students, J.L.D. Godfrey collected at BridalVeil Falls for the preparation of a flora of the hepatics and hornworts of southwestern BritishColumbia (Godfrey 1977). Another student of W.B. Schofield, J.P. van Velzen, also sampledat the site for his work on epilithic bryophyte communities in southwestern British Columbia(Van Velzen 1981). D.H. Vitt also did some bryological work at Bridal Veil Falls, collectingspecimens for a study of foliicolous bryophytes and lichens of Thuja plicata in westernBritish Columbia (Vitt et al. 1973). Four foliicolous species of Orthotrichum were foundgrowing in abundance at Bridal Veil Falls, the first report of foliicolous mosses in NorthAmerica north of the tropics. Two species of bryophytes rare to British Columbia,Hymenostylium insigne and Barbula amplemfolia were first collected at Bridal Veil Falls(Zander and Eckel 1982). The collection of Hymenostylium insigne by W.B. Schofield wasthe first collection of this species in the New World. Seligeria tristichoides was also firstcollected in British Columbia on the cliffs at Bridal Veil Falls by F.J. Hermann (Crum 1975),the first report of this species in western North America.16Chapter 2. Inventory and keys2.1. Materials and methods2.1.1. CollectionA list of the bryophytes found at Bridal Veil Falls was compiled primarily fromcollections made at the site during the course of this study. Field work was conducted during1991 and 1992. Of 661 specimens collected, a total of 996 bryophytes were identified. Forcompleteness, I also examined specimens collected at Bridal Veil Falls by earlier bryologistsand deposited at the herbarium of the University of British Columbia (UBC). These wereincluded in the flora.Field collections were made from as many types of microhabitats as could berecognized, taking into account accessibility and time constraints. Each specimen was given acollection number and ecological information and date of collection were recorded. Ecologicaldata recorded included type of habitat and substratum, moisture and light conditions, andexposure (compass direction faced), when applicable.2.1.2. Identification and preparation of the keysFor the identification of the mosses collected during this study, I referred primarily tothe Moss flora of the Pacific Northwest by E. Lawton (1971). Keys to the mosses of BridalVeil Falls were prepared based mainly on personal observations made in the process ofidentification. In the case of species that were not collected during the course of this study, Iexamined herbarium specimens collected by other bryologists at Bridal Veil Falls. The work17of the following authors guided my observations and provided useful characters to be used inthe construction of the keys:Crum and Anderson (1981);Dixon and Jameson (1924);Frisvoll (1983, 1985, 1988);Gangulee (1972);Hill (1984);Hoisington (1979);Ireland (1969a, 1969b, 1970, 1976, 1982);Jamieson (1976);Koponen (1974);Lawton (1965, 1967, 1971);Nyholm (1975);Schofield (1969b);Schofield and Talbot (1991);Shaw (1982);Smith (1978);Vitt (1973);Vitt and Buck (1992);Vitt et al. (1988);Zander (1978, 1979);Zander and Eckel (1982).For the identification of the liverworts, I used the flora of the hepatics and homwortsof southwestern British Columbia prepared by Godfrey (1977). Much information on theliverwort flora of Bridal Veil Falls can be derived from Godfrey's work, as all the species ofliverworts found are treated in her keys and descriptions. I also used the following floras andtaxonomic treatments which provided helpful additional descriptions and illustrations:Frye and Clark (1937-1947);Hicks (1992);Macvicar (1960);Schuster (1966, 1969, 1974, 1980, 1992a, 1992b);Smith (1990).A set of voucher specimens is deposited in the herbarium of the University of BritishColumbia (UBC). UBC accession numbers for these specimens are given in Appendices Aand B. In most cases, voucher specimens were collected during the course of this study. Theexception is for species not found during this study, but collected by others. In such cases,carefully examined herbarium specimens serve as vouchers.182.1.3. NomenclatureThe nomenclature used in this study follows current checklists of the mosses andliverworts of North America (Stotler and Crandall-Stotler 1977; Anderson 1990; Anderson etal. 1990). My interpretation of certain species however, occasionally differed from that of theauthors of these checklists In these particular cases, the following nomenclature was used:a) Mosses:-Barbula coreensis (Card.) Saito is treated as a synonym of Barbula amplexifolia(Mitt.) Jaeg. based on Zander (1979);- Brachythecium frigidum (C.Muell.) Besch. and Brachythecium asperrimum (Mitt.)Sull. are considered distinct species based on the work of Hoisington (1979);-Ditrichum flexicaule (Schwaegr.) Hampe and Ditrichum crispatissimum (C.Muell.)Par. are considered distinct species based on Frisvoll (1985);- Schistidium apocarpum var. strictum (Turn.) Moore is treated as distinct from S.apocarpum var. apocarpum (Hedw.) Bruch & Schimp. in B.S.G. as described in Lawton(1971).b) Liverworts:-Chiloscyphus pallescens (Ehrh. ex Hoffm.) Dum. is considered as a synonym ofChiloscyphus polyanthos (L.) Corda based on Godfrey (1977) and Smith (1990);-Plagiochila asplenioides (L.) Dum., P. porelloides (Torrey ex Nees) Lindenb. and P.satoi Hatt. are treated together as P. asplenioides sensu law based on Godfrey (1977) (seeSection 2.2.3 for discussion).192.2. Results and discussion2.2.1. Diversity of the floraThe known bryophyte flora of Bridal Veil Falls Provincial Park and adjacent boulderslope area was found to be composed of 210 species. This includes 150 species of mosses andtwo additional varieties in 86 genera, and 60 species of liverworts in 36 genera. It represents22% of the moss flora of the province and 27% of its liverwort flora (data from Schofield1968a and Ireland et al. 1987). The bryophytes known from Bridal Veil Falls are listed inAppendices A and B.Prior to this study, only 184 species were known for the study area. Species added tothe flora by virtue of this study are indicated in the annotated list of species (Appendices Aand B). A variety of Barbula convoluta, B. convoluta var. gallinula Zander, was found to benew to British Columbia (see Section 2.2.4). Two other species, Hymenostylium insigne andAnastrophyllum assimile are known from only very few other localities in British Columbia.Finally, Barbula amplexifolia is known in British Columbia only from Bridal Veil Falls.The number of bryophyte species at Bridal Veil Falls is relatively high compared toplaces of similar area in the Lower Mainland of British Columbia. Lynn Canyon Park (25 ha)in North Vancouver, for instance, has a bryophyte flora composed of 100 species of mosses(Krause and Schofield 1977) and approximately 54 species of liverworts (W.B. Schofield,pers. communication). Similarly, the bryoflora of Pacific Spirit Park (763 ha) in Vancouverconsists of 122 species of mosses and 38 species of hepatics (Schofield 1992, unpublisheddata). Of the 166 species composing the hepatic flora of Southwestern British Columbia(Godfrey 1977), 36% are present at Bridal Veil Falls; 56% of the 64 genera described byGodfrey (1977) are represented.202.2.2. Keys to the mosses of Bridal Veil Falls Provincial Park and adjacent boulderslope.The following dichotomous keys treat the moss flora of Bridal Veil Falls ProvincialPark and adjacent boulder slope area. They should be used in conjunction with Lawton'sMoss Flora of the Pacific Northwest (1971) which provides illustrations and descriptions ofmost of the species treated here. Her flora also provides a glossary of specialized terminologyused throughout the following keys. A list of synonyms for the species named differently thanin Lawton or references for species not treated in Lawton is provided in Appendix D.The first key leads to genera. Genera can then be keyed to species in a following setof keys. The exceptions are unispecific genera or isolated species within certain genera inwhich case species is given in the first key. In some cases, several different routes will leadto the same species. This was done to accommodate possible differences in interpretation bydifferent observers.The keys are based primarily on gametophytic characters. Sporophytic characters areused only when providing determinant additional information or in the case of species whichare otherwise difficult or impossible to distinguish. Most characters are microscopic requiringthe use of dissecting and compound microscopes.21Keys to the mosses of Bridal Veil Falls and adjacent boulder slope1-General key1.^Leafy plants not apparent, sporophyte solitary; seta stout, capsule inclined with theupper face flattened; plant small. ^  Buxbaumia pipen1. Leafy plant apparent, with or without sporophyte ^  22. Leaf cells arranged in a reticulate pattern with narrow, elongate, chlorophyllous cellssurrounding large, swollen, porose, fibrillose, dead, hyaline cells; branches in fascicles.^  Sphagnum (p. 48)2.^Leaf cells mostly chlorophyllous (with the exception of marginal, alar, or basal cells);branching uncommon, irregular, pinnate, or dendroid. ^  33.^Gemmae lens-shaped, contained in terminal gemmae cups; peristome of four, rigidteeth. ^  Tetraphis (p. 48)3. Gemmae, if present, never in gemmae cups, or gemmae absent; peristome absent, or ofmore than four teeth, or sporophyte absent. ^  44. Leaves with lamellae on the upper surface.  54.^Leaves without lamellae on the upper surface. ^  95.^Leaves bordered by elongate cells, undulate; lamellae few, restricted to the uppersurface of the costa. ^  Atrichum (p. 38)5. Leaves not bordered, not undulate; lamellae usually numerous, not restricted to theupper surface of the costa. ^  66. Lamellae found on the lower surface of the leaf, and on the upper surface of the costa.^  Oligotri chum aligerum6.^Lamellae restricted to the upper surface of the leaf. ^  7^7.^Apical cells of the lamellae smooth ^  Polytrichum (p. 45)7.^Apical cells of the lamellae papillose  822^8.^Apical cells of the lamellae longer than broad in cross-section, somewhat ovate, withthe outer wall strongly thickened; leaves somewhat flexuose when dry. Polytrichastrum alpinum8. Apical cells of the lamellae not longer than broad in cross-section, rounded at the top,with the walls more or less evenly thickened; leaves stiff and erect when dry.^  Pogonatum urnigerum9. Leaves bordered by a band of narrowly elongate cells sharply contrasting with thecells within; leaf cells smooth. ^  109.^Leaves not bordered, marginal cells not strongly differentiated from the rest of thelamina, or cells gradually longer, and narrower toward the margin; leaf cells smooth orpapillose. ^  1510.^Leaf bases split in two flaps at the base, clasping the stem and the leaf above; leavescomplanate, distichous; plant small. ^  Fissidens bryoides10. Leaf bases not of two flaps, not clasping the stem ^  1111. Leaf margins recurved, and unistratose ^  Blyum (p. 39)11.^Leaf margins plane, unistratose or multistratose. ^  1212.^Leaf margins entire, and multistratose. ^ Rhizomnium glabrescens12. Leaf margins toothed, or if entire, then unistratose. ^  1313. Costa extending to about 3/4 the length of the leaf; leaf cells elongate, 2:1 or longer;leaf margins unistratose; plant small and delicate. ^ Epipterygium tozeri^13.^Costa extending to near apex, percurrent or excurrent; leaf cells (with exception ofmarginal cells) short, usually less than 2:1. ^  1414.^Leaf margins multistratose, marginal teeth double ^ Mnium (p. 43)14. Leaf margins unistratose, marginal teeth single, or absent. . . .^Plagiomnium (p. 44)15. Leaves complanate (flattened) on stem ^  1615.^Leaves not complanate on stem, regularly distributed around the stem ^ 2716.^Leaves arranged in two rows. ^  1716.^Leaves arranged in more than two rows. ^  1923^17.^Leaves with a long, narrow, papillose awn; leaf bases sheathing the stem; plant greenabove, brown below. ^  Distichium capillaceum17. Leaves not awned.  1818. Costa extending to about mid-leaf; leaves spatulate, widest above middle, apex broadlyrounded; leaf bases not of two flaps. ^ Homalia trichomanoides^18.^Costa extending well beyond mid-leaf to apex or just below; leaves not spatulate,widest at or below the middle, apex acute or obtuse; leaf bases split in two flaps at thebase, clasping the stem and the leaf above. ^ Fissidens (p. 42)19.^Costa single, to mid-leaf or beyond.  2019. Costa absent, or short, or short and double. ^  2220. Stem with abundant paraphyllia; leaves undulate, lingulate, apex rounded, abruptlyending in a short point; capsule immersed; flagelliform branches often present.^  Metaneckera menziesii^20.^Stem without, or with occasional paraphyllia; leaves not undulate; capsule exserted. 2121.^Median leaf cells short to broadly elongate, usually less than 5:1   2221. Median leaf cells narrowly elongate, more than 5:1. ^  2722. Leaves spatulate, widest above the middle, apex broadly rounded; plant stronglycomplanate, and glossy. ^  Homalia trichomanoides^22.^Leaves narrower above than at mid-leaf, apex acute or obtuse; plant more or lesscomplanate, dull or glossy. ^  Porotrichum (p. 46)23.^Plant with tiny, filiform, branch-like propagula clustered in leaf axils. Pseudotaxiphyllum elegans23. Plant without asexual reproductive bodies, or if present, not branch-like. ^ 2424. Leaves decurrent ^  Plagiothecium (p. 45)24.^Leaves not decurrent.  2525.^Alar cells differentiated, forming a distinct group of swollen cells. . Hypnum lindbergii^25.^Mar cells not differentiated, or if differentiated then not swollen ^ 262426.^Leaves ovate-lanceolate, apex gradually narrowing into a point. Pseudotaxiphyllum elegans26. Leaf ungulate to ovate-lanceolate, more or less rounded below the pointed apex, oftenundulate. ^  Neckera (p. 43)27. Leaf cells conspicuously sinuose; leaves costate, usually imbricate when dry (with theexception of hair-points, if present, which are often squarrose or reflexed, and more orless twisted). ^  2827.^Leaf cells not sinuose, or leaves without a costa, or crisped and contorted when dry.3128.^Leaf cells multipapillose, or leaves muticous and rounded at apex, or hair-pointspapillose ^  Racomitrium (p. 46)28. Leaf cells smooth, or with papillae on the back of the costa and scattered sparsely onthe lamina in the upper leaf; leaves acute at apex, hyaline hair-points, if present,denticulate but not papillose. ^  2929. Cells at mid-leaf elongate, more than 2:1, commonly 3-4:1. . ^Racomitrium (p. 46)29.^Cells at mid-leaf short, 1-2:1. ^  3030.^Plant dark green to nearly black; leaf cells smooth. Schistidium apocarpum var. apocarpum30. Plant often reddish brown; leaf cells at the back of the costa and upper laminapapillose. ^  Schistidium apocarpum var. strictum31. Plants minute, up to 4 mm including the sporophyte (gametophyte 1 or 2 mm high);leaves costate, lanceolate, with smooth cells. ^ Seligeria (p. 47)31.^Plants larger, or if not, leaves ecostate, or leaf cells multipapillose   3232.^Leaf cells papillose ^  3332. Leaf cells smooth.  6533. Stems papillose ^  3433.^Stems smooth.  352534.^Leaves ovate-lanceolate, somewhat cordate; costa obscure, usually forked, ending at orbefore mid-leaf; plant with many lateral branches at right angles with the main branch. Heterocladium macounii34. Leaves linear to lanceolate; costa strong, single, ending just before apex; plantunbranched, or with few irregular, erect branches. . .. Hymenostylium recurvirostre35. Leaves without a costa, or with a short single costa ending before mid-leaf, or with adouble costa ^  3635.^Leaves with a well-defined single costa ending above mid-leaf. ^ 4136.^Leaves with a long double costa extending to mid-leaf or beyond; leaves plicate. Rhytidiadelphus triquetrus36. Leaves without a costa, or costa single, or if double, not extending beyond mid-leaf.3737. Stems and branches with abundant paraphyllia; plant much branched, bi- to tripinnate,frond-like; new growth arising from the middle of previous year's stem.^  Hylocomium splendens37.^Stems and branches without paraphyllia, or if paraphyllia present, plant irregularlybranched ^  3838.^Leaves ending in a whitish hair-point. ^  3938. Leaves lacking a whitish hair-point.  4039. Leaf cells with forked papillae; capsule immersed. ^ Hedwigia ciliata39.^Leaf cells with simple papillae; capsule exserted. ^ Pseudobraunia cahfornica40.^Costa single, often forked, obscure; leaves with 2 or more papillae per cell; stempapillose. ^  Heterocladium macounii40. Costa double, or absent; leaves with one papilla per cell, papillae formed by projectingupper ends of cells; stem smooth. ^ Pterigynandrum fihforme41. Plants with numerous branches, forming a mat over the substratum or dendroid. . 4241.^Plants erect, scarcely branched, usually in tufts ^  452642.^Stems with numerous paraphyllia; leaf cells with 1 papilla per cell located at the upperend of the cells; upper leaves without hair-points ^  4342. Stems without paraphyllia, or paraphyllia scarce; leaf cells multipapillose, orunipapillose with the papillae not associated with the upper end of the cell; upperleaves usually with hair-points ^  4443. Plants dendroid; branch leaves obtuse and serrate at apex; costa rough with projectionson the dorsal surface; branches circinate when dry. ^ Dendroalsia abietina43.^Plants forming a mat over the substratum; branch leaves entire, or slightly serrulateabove; costa smooth ^  Pseudoleskea stenophylla44.^Leaves imbricate when dry, margins recurved; papillae scarce, restricted to costa andcells in upper leaf; costa percurrent; plants dark green to reddish brown. Schistidium apocarpum var. strictum44. Leaves crisped and contorted when dry, margins plane; papillae abundant, found oncells throughout most of the leaf; costa ending before apex, pellucid; plants green toyellow-green. ^  Claopodium (p. 40)45. Leaf margins coarsely serrate near apex. ^  4645.^Leaf margins entire or serrulate, or if toothed, the teeth very distant, or few andrestricted to the very tip of the leaf. ^  5046.^Leaf cells with numerous cuticular thickenings resembling papillae; leaf apices rough,with teeth on the margins, on the back of the costa, and often on the lamina. Plagiopus oederiana46. Leaf cells mammillose (bulging), or with one papilla per cell. ^ 4747. Plant with clusters of orange-brown rhizoids matting the base of the stem, dull,yellowish-green; leaf margins bistratose above, teeth often double; capsule green,globulose when young, ribbed when dry. ^ Bartramia pomiformis47.^Plant without clusters of rhizoids matting the base of the stem, or if so, plants not dull,but somewhat shiny; leaf margins unistratose, teeth single ^ 4848.^Leaves linear-lanceolate from an orange-brown, sheathing leaf base, apex acute. Timmia austriaca48.^Leaves long lingulate, ovate-lanceolate, or lanceolate, base not sheathing, apex acute torounded. ^  492749.^Leaf cells mammillose (bulging) on one or both surfaces; leaves crisped and contortedwhen dry. ^  Dichodontium pellucidum49. Leaf cells with one large papilla on each surface; leaves imbricate-flexuose when dry;gemmae frequent, borne in a globular head at the tip of a leafless stalk at the tip ofthe shoot. ^  A ulacomnium androgynum50. Plants with ovoid, multicellular, often massive gemmae borne in leaf axils; leavesovate to ovate-lanceolate; leaf cells multipapillose, short above, short rectangularbelow; plants small.^  Barbula (p. 38)50.^Plants without gemmae, or gemmae otherwise; leaves linear, lanceolate, ovate-lanceolate, oblong, lingulate or spatulate. ^  5151.^Leaves broad throughout, oblong, ungulate, or spatulate, apex usually rounded tobroadly pointed; costa often excurrent in a stiff point or awn; upper leaf cells shortand multipapillose, lower leaf cells elongate and smooth ^ 5251. Leaves narrowed toward apex, apex acute or narrowly obtuse, or leaf cells with only 1papilla per cell on each surface of the leaf. ^  5552. Plants with abundant gemmae clustered in leaf axils, gemmae consisting of branched,uniseriate filaments. ^  Encalypta procera52.^Plants without gemmae.  5353.^Leaf margins differentiated, cells longer and narrower than within, smooth or with lowpapillae; costa excunent as a coloured point or awn. ^ Tortula subulata53. Leaf margins not differentiated; costa variable.  5454. Calyptra long campanulate; peristome single or double, teeth not twisted; basal leafcells (best observed near the base of the costa) with thickened, coloured crosswalls;costa ending before apex, percuffent, or excurrent as a short point not readily visibleunder low magnification (10x), or if so, the long hair-point hyaline, smooth, andflexuose ^  Encalypta (p. 41)54.^Calyptra cucullate; peristome single, teeth long, fused below, and twisted together;basal leaf cells with evenly thickened crosswalls; costa excurrent as a long, toothedawn readily visible at low magnification (10x). ^ Tortula (p. 48)55.^Leaves (at least the upper ones) with hyaline, denticulate hair-points; plants dark greento reddish brown ^  Schistidium apocarpum var. strictum55.^Leaves without hyaline hair-points, sometimes apiculate, apiculus never denticulate.562856.^Gemmae usually abundant, borne in leaf axils near the tip of the stem, consisting of 3-to 4-celled elliptic bodies; leaf cells multipapillose, papillae more than 2 per cell. Zygodon viridissimus56. Gemmae absent, or if present leaf cells with 1-2 or no papillae. ^ 5757. Leaves ovate-lanceolate, broadened at the base. ^  5857.^Leaves linear to lanceolate, not broadened at the base.  5958.^Plants usually on trees; basal leaf cells usually very thick-walled (often as thick as thecells are wide) and coloured, often pitted; calyptra hairy and campanulate. Ulota (p. 48)58. Plants usually on rock or soil, occasionally on logs; basal leaf cells usually hyalineand not pitted; calyptra smooth and cucullate. ^ Didymodon (p. 41)59. Basal leaf cells hyaline, forming a "V" at the base of the leaf (i.e. hyaline cellsextending higher along the margins of the leaf than along the costa) ^^  TorteIla tortuosa59.^Basal leaf cells not hyaline, or if hyaline, not arranged in a conspicuous "V". . .. 6060.^Mar cells differentiated, forming a large group of swollen cells; leaves usually falcate-secund, costa long excurrent. ^  Dicranum fuscescens60. Mar cells not differentiated, or not inflated; leaves straight, costa ending before apexto shortly excurrent. ^  6161. Leaf cells with 1 or 2 papillae on each surface; plants usually on trees.^  Orthotrichum (p. 44)61.^Leaf cells multipapillose, with more than 2 papillae on each surface; plants usually onrock. ^  6262.^Costa often shortly excunent as a smooth terminal point, and extreme tip of the leavesoften with a few smooth teeth; capsule wrinkled when dry, peristome single, of 16teeth; plants green above, reddish brown below, leaf bases sheathing the stem.^  Bryoerythrophyllum recurvirostre62.^Costa usually ending below apex, or if excurrent in a smooth terminal point, then leafapex not denticulate; capsule ribbed or smooth when dry, peristome absent . .. . 632963.^Leaves usually tightly twisted and contorted when dry; upper basal leaf cells withnumerous oval papillae resembling cuticular ridges; papillae often found on thecrosswalls of leaf cells; capsule ribbed when dry. ^ Amphidium (p. 38)63. Leaves usually loosely twisted and contorted when dry, or somewhat imbricate; upperbasal leaf cells smooth, or with a few, rounded papillae similar to the papillae on theupper leaf cells; papillae not on the crosswalls of the leaf cells; capsule smooth whendry.   6464. Leaf margins usually plane and entire, or often crenulate along the upper basal regionwhere the leaves curve back; capsule pale brown, reddish at the mouth; operculum notattached to the columella. ^  Gymnostomum aeruginosum64.^Leaf margins usually recurved at middle on one or both sides, entire; capsule darkbrown; operculum often attached to the columella. ^ Hymenostylium (p. 42)65.^Costa absent, or double, or short, ending before mid-leaf.  6665. Costa single (occasionally forked), extending to mid-leaf or beyond. ^ 7866. Stems with abundant paraphyllia; stem leaves with a long, attenuate, twisted apex;branching regularly bi- to tripinnate; new growth arising from the middle of theprevious year's stem ^  Hylocomium splendens66.^Stems without paraphyllia, or paraphyllia scarce; stem leaves not twisted at apex;branching irregular, or mostly once pinnate. ^  6767.^Leaves plicate. ^  Rhytidiadelphus (p. 46)67. Leaves smooth, not plicate. ^  6868. Alar cells with a distinctly differentiated single row of 3-4 elongated and inflated cells.^  Brotherella roellii68.^Mar cells not differentiated, or if differentiated, never with a single row of 3-4elongated and inflated cells. ^  6969.^Cortical cells of stem enlarged and thin-walled; alar cells inflated. ^ 7069. Cortical cells of stem small and thick-walled; alar cells not inflated, or if so, leavessquarrose. ^  7170. Leaves obtuse to rounded at apex, straight. ^ Calliergonella cuspidata70.^Leaves acute at apex, usually falcate-secund.  Hypnum lindbergii3071.^Stem red; alar cells differentiated, smaller, thick-walled and coloured (orange-brown). Pleurozium schreberi71. Stem green or brown. ^  7272. Leaves narrowly acuminate from a cordate base, squarrose or wide-spreading (upper,lanceolate portion of the leaf pointing away from the stem). ^ 7372.^Leaves ovate-lanceolate to lanceolate, imbricate to spreading, or leaves falcate. . . 7473.^Mar cells clearly differentiated, short rectangular to rounded, or enlarged and inflated;leaves often shortly decurrent. ^  Campylium stellatum73. Mar cells not clearly differentiated, hyaline, shorter and wider than surrounding cells;leaves not decurrent ^  Rhytidiadelphus squarrosus74. Leaves concave, entire, or serrate to serrulate at apex. ^  7574.^Leaves not concave, entire, or serrulate at leaf base and occasionally at apex. . ^ 7675.^Leaves decurrent, decunent portion narrow, of elongate, hyaline cells tapering to thestem (often remaining on the stem); leaf apices often recurved. Plagiothecium cavifolium75. Leaves not decurrent; leaf apices not usually recurved. ^ Hygrohypnum luridum76. Leaves falcate to circinate, slenderly acuminate to a filiform point; plants irregularly ormore or less regularly pinnate, branches numerous. ^ Hypnum (p. 43)76.^Leaves straight to somewhat falcate-secund, never circinate; plants irregularlybranched, branches few. ^  7777.^Leaves less than 0.5 mm long; plants minute, filiform. Platydictya jungermannioides77. Leaves more than 0.5 mm long; plants short, thin. ^ Isopterygiopsis pulchella78. Leaves lanceolate from an orange-brown sheathing leaf base; leaf cells in cross-sectionbulging on the upper surface of the leaf. ^ Timmia austriaca78.^Leaves not lanceolate, or leaf bases not sheathing, or if sheathing, not coloured. . 7979.^Plants dendroid, branches clustered at the top of a usually erect stem ^ 8079.^Plants not branching, or branching irregular or pinnate ^  813180.^Branch leaves decuffent; alar cells not differentiated. . .. Leucolepis acanthoneuron80. Branch leaves not decurrent, rounded at insertion, forming auricles just above the alarregion; alar cells coloured and swollen. ^ Climacium dendroides81. Leaves ovate, strongly concave, imbricate, barely changed when dry; costa strong,ending just before apex, percuffent or excunent. ^  8281.^Leaves lanceolate, or not concave or imbricate, or if so, costa shorter, ending wellbefore leaf apex. ^  8482.^Leaves apiculate; plants whitish green above, pinkish below. . . . . Plagiobryum zierii82. Leaves rounded to broadly acute, without an apiculus. ^  8383. Plant yellow-green. ^  Anomobryum fihforme83.^Plant with a reddish wine tinge. ^  Bryum miniatum84.^Alar cells differentiated, forming a distinct group of inflated cells. ^ 8584. Alar cells not differentiated, or if differentiated, not forming a distinct group ofinflated cells. ^  9185. Leaves plicate, falcate-secund. ^  8685.^Leaves not plicate, or leaves straight.  8786.^Stem with abundant paraphyllia; alar cells often coloured. . . Palustriella commutata86. Stem without paraphyllia (pseudoparaphyllia sometimes present); alar cells hyaline.^  Sanionia uncinata87. Alar cells coloured; costa strong, often excurrent; leaves narrowly ovate to lanceolate.8887.^Alar cells hyaline, or if coloured, costa never excurrent; leaves ovate to ovate-lanceolate. ^  8988.^Costa excurrent from an ovate leaf base. ^  Blindia acuta88.^Costa excurrent or ending before apex, leaves long, lanceolate. . ^Dicranum (p. 40)3289.^Median leaf cells mostly shorter than 5:1; costa extending well into leaf apex. Cratoneuron filicinum89. Median leaf cells mostly longer than 5:1; costa not extending to leaf apex. ^ 9090. Leaves straight to often falcate, acute or obtuse, not decurrent, smooth, margin entire.^  Hygrohypnum luridum90.^Leaves straight, acute, usually decurrent at the base, plicate or smooth, margin usuallyserrulate nearly to the base, or entire. ^ Brachythecium (p. 39)91.^Stem with abundant paraphyllia; plants usually on trees. ^ 9291. Stem without abundant paraphyllia or paraphyllia scarce.  9392. Leaves rounded at apex. ^  Metaneckera menziesii92.^Leaves gradually acuminate.  Pseudoleskea stenophylla93.^Plant blackish or brownish, erect, scarcely branched; leaves often with hair-points; leafcells thick-walled. ^  9493. Plant yellow-green, green or dark green (sometimes reddish), erect or prostrate,branching scarce, or pinnate, or irregular; leaves without hair points ^ 9794. Cells in upper part of the leaf elongate. ^  Pohlia nutans94.^Cells in upper part of the leaf short, somewhat quadrate or rounded. ^ 9595.^Leaves tightly crisped and contorted when dry; plants in dense tufts. Grimmia torquata95. Leaves imbricate or loosely contorted when dry. ^  9696. Hair-points present on upper leaves. ^ Schistidium apocarpum var. apocarpum96.^Hair-points absent. ^  Didymodon rigidulus var. gracilis97.^Plants usually prostrate (sometimes erect), forming mats over the substratum;branching irregular or pinnate. ^  9897. Plants usually erect, forming tufts; branching absent or scarce. ^ 11698. Leaf margins recurved from base to near apex. ^  9998.^Leaf margins plane or recurved at the base only.  1003399.^Leaves plicate, not concave; costa single. ^ Homalothecium (p. 42)99. Leaves not plicate, concave; costa often with supplementary costae flaring at its base.^  Antitrichia (p. 38)100. Branch leaves squarrose, entire or minutely serrulate at the base. . Campylium (p. 40)100. Branch leaves erect or spreading, not squarrose, or if so, leaves serrate. ^ 101101. Leaves plicate ^  102101. Leaves not plicate  104102. Leaf margins often recurved, serrate or toothed at the apex and often at the base.^  Homalothecium (p. 42)102. Leaves margins plane or recurved at the very base only, entire to serrulate or serrate.^  103103. Alar cells more or less inflated; operculum conic. ^ Brachythecium (p. 39)103. Mar cells not inflated; operculum long rostrate. ^ Eurhynchium (p. 41)104. Median leaf cells short, 1-2(3):1; leaves ovate to ungulate, with a broad apex. . 105104. Median leaf cells long and narrow, more than 3:1, or if shorter, leaves ovate-lanceolatewith an acuminate apex. ^  106105. Costa strong, ending before apex, toothed at the back near apex; leaves coarselyirregularly toothed at apex; plant coarse ^ Thamnobryum neckeroides105. Costa short, extending to the middle or 3/4 of the leaf, not toothed at the back; leavesserrate to serrulate at apex; plant small, in thin mats. . . Porotrichum vancouveriense106. Leaf margins serrate or serrulate from apex to nearly base. ^ 107106. Leaf margins entire or serrulate above (occasionally to below mid-leaf, but never tobase). ^  111107. Leaf margins irregularly toothed; alar cells differentiated, forming a distinct, smallgroup of dense and thick-walled, quadrate, rounded or irregular cells.^  Isothecium myosuroides107. Leaf margins regularly toothed. ^  10834108. Leaf apices broadly acute to rounded; cells at the extreme apex of the leaves usuallyshort, 2:1 or less. ^  109108. Leaf apices acute to acuminate, never obtuse or rounded; cells at the extreme apex ofthe leaves elongate, mostly longer than 2:1. ^  110109. Mar cells not clearly differentiated, elongate, wider than other cells, occasionallyslightly inflated. ^  Platyhypnidium riparioides109. Mar cells differentiated, short, quadrate, rounded or irregular.^ Eurhynchium pulchellum (see generic key for variety, p. 41)110. Stem and branch leaves different in form (branch leaves ovate, stem leaves larger,cordate, decurrent, and often squarrose); operculum long rostrate.^  Eurhynchium (p. 41)110. Stem and branch leaves not different in form; operculum conic.^Brachythecium (p. 41)111. Leaves narrowly ovate -lanceolate, tapering to a long, channelled (margins rolledinward) apex. ^  Campylium polyganum111. Leaves ovate to ovate-lanceolate, apex not long and channelled. ^ 112112. Leaves concave. ^  113112. Leaves not concave.  115113. Costa single, often ending in a tooth at the back (look at several leaves).^  Scleropodium (p. 47)113. Costa single or often double or forked, not ending in a tooth ^ 114114. Leaves often falcate, entire at apex. ^ Hygrohypnum luridum114. Leaves straight, usually finely serrulate at apex. ^ Brachythecium (p. 39)115. Leaves small, mostly less than 1 mm long, not plicate or decurrent; alar cells notinflated. ^  Amblystegium serpens115. Leaves larger, mostly more than 1 mm long, often plicate and more or less decurrent;alar cells usually more or less inflated. ^ Brachythecium (p. 39)116. Leaf margins involute, at least above.  117116. Leaf margins plane or recurved. ^  11935117. Leaves erect, narrowly lanceolate, gradually tapering from base to apex.^  Ditrichum crispatissimum117. Leaves spreading or squarrose, lanceolate from a broadened, sheathing leaf base. . 118118. Capsules long and narrow, cylindric; stems rarely taller than 5 mm.^  Trichodon cylindricus118. Capsules short, zygomorphic; stems up to 30 mm. ^ Dicranella schreberiana119. Leaf margins toothed above. ^  120119. Leaf margins entire or serrulate above ^  124120. Leaf cells elongate, more or less uniform throughout the length of the leaf.^  Pohlia (p. 45)120. Leaf cells short above and elongate below, or vice-versa. ^ 121121. Leaf cells elongate above, quadrate at the base. ^ Anacolia menziesii121. Leaf cells quadrate above, elongate at the base.  122122. Leaf margins bistratose above, teeth often double ^ Bartramia pomiformis122. Leaf margins unistratose throughout, teeth single.  123123. Leaf apices rough with teeth on the margins, on the back of the costa, and often onthe lamina; stem with clusters of reddish-brown rhizoids. . . . . Plagiopus oederiana123. Leaf apices with teeth on the margin only, and papillae on the back of the costa; stemwithout clusters of rhizoids. ^  Dichodontium pellucidum124. Leaf cells thick-walled, short, uniform throughout the length of the leaf.^  Tetraphis (p. 48)124. Leaf cells thick- or thin-walled, elongate throughout or short above to graduallyelongate below. ^  125125. Leaves abruptly lanceolate from a broad apex. ^  126125. Leaves linear, ungulate, ovate, or lanceolate, gradually tapering towards apex. ^ 128126. Leaves crisped and contorted when dry; plants on trees. ^ Ulota126. Leaves spreading and loosely contorted when dry; plants on soil. ^ 12736127. Capsules long and narrow, more than 4X longer than wide, cylindric; stem rarelyhigher than 5 mm. ^  Trichodon cylindricus127. Capsules short, less than 3X longer than wide, zygomorphic; stem up to 30 mm.^  Dicranella schreberiana128. Leaves cells elongate throughout, thin-walled, gradually longer and narrower towardsthe margin. ^  129128. Leaves cells short, or if elongate, cells somewhat thick-walled, or not graduallydifferentiated towards the margin ^  130129. Costa extending to 3/4 the length of the leaf or above; stem usually more than 1 cmlong ^  Pohlia (p. 45)129. Costa not extending above 3/4 the length of the leaf; stem usually less than 1 cm long.^  Epipterygium tozeri130. Leaves crisped and contorted when dry. ^  131130. Leaves erect or spreading when dry, sometimes slightly contorted. ^ 132131. Leaf margins recurved from near base to near apex; leaf margins bistratose above.^  Dicranoweisia cirrata131. Leaf margins plane or more or less recurved below; leaf margins unistratose.^  Dichodontium pellucidum132. Costa excurrent as a short awn ^  Pottia truncata132. Costa ending before apex, percurrent or shortly excurrent, but never as an awn. ^133133. Leaf cells short and quadrate above, gradually longer below. . . Ceratodon purpureus133. Leaf cells more or less elongate throughout. ^  134134. Leaf margins bistratose above. ^  Dicranella varia134. Leaf margins unistratose.  135135. Median leaf cells narrow, less than 20 gm wide; capsule symmetric. . Pohlia nutans135. Median leaf cells broad, more than 20 gm wide; capsule zygomorphic.^  Funaria hygrometrica372-Keys to multispecific genera.Amphidium1.^Leaf margins plane (or only slightly recurved below) and more or less entire; papillaeon leaf cells round and wide. ^  A. lapponicum1.^Leaf margins usually recurved and often with distant, blunt teeth above; papillae onleaf cells elliptical, small, resembling cuticular ridges. ^ A. californicumAntitrichia1.^Supplementary costae strong; leaf cells pitted; leaves loosely imbricate when dry, tipsoften spreading. ^  A. curtipendula1.^Supplementary costae weak or absent; leaf cells not pitted, or very weakly so; leavestightly imbricate when dry. ^  A. californicaAtrichum1.^Monoicous. ^  A. undulatum1. Dioicous.  22. Leaf cells more or less rounded with pronounced corner thickenings; dioicous ormonoicous. ^  A. undulatum2.^Leaf cells more or less angular, most corner thickenings not well defined, walls moreor less evenly thickened; always dioicous. ^  A. selwyniiBarbula1.^Leaves rounded at apex, not sheathing at the base; costa ending a few cells beforeapex; gemmae usually larger than 100 pm ^ B. convoluta var. gallinula1.^Leaves acute at apex, somewhat sheathing at the base; costa excurrent in a short point;gemmae usually 100 Inn or shorter. ^  B. amplexifolia38Brachythecium1.^Leaves concave, tapering more or less abruptly to a short point. ^ 21. Leaves not concave, usually narrowly acuminate. ^  42. Alar cells short-rectangular or quadrate, not inflated and often dense and more or lessopaque. ^  B plumosum2.^Mar cells rectangular or hexagonal, more or less inflated and hyaline. ^ 33.^Mar cells forming distinct decurrent auricles of inflated cells, sharply set-off, easilydistinguished at low magnification (20X on the dissecting microscope). . . . B. rivulare3. Mar cells not forming distinct decunent auricles, more or less inflated, not sharply set-off. ^  B. rutabulum4. Leaves straight to falcate, sometimes asymmetric, usually not plicate, narrowly ovatebelow, gradually tapering to a long, narrowly acuminate apex; alar cells few, quadrate,irregular or short rectangular, not swollen. ^  B. velutinum4.^Leaves straight, symmetric, usually plicate, broad below, narrowly attenuate at apex. 55.^Stem leaves deeply plicate, most with at least two laminal (not marginal) plications oneither side of the costa. ^  B. frigidum5. Stem leaves plane, wrinIded or with only one plication on either side of the costa. ^66. Inflated ajar cells in a well defined, often decurrent, group of ten or more oval orround cells; inflated cells plainly visible under low magnification (40X). . B. frigidum6.^Enlarged alar cells less numerous, (usually 6-9), these being more oblong orrectangular in outline rather than round, not plainly visible under low magnification(40X). ^  B. asperrimumBryum1.^Marginal leaf cells not differentiated; leaves rounded at apex, margins plane; plantwith a wine red tinge, leaves imbricate when dry. ^ B. miniatum1.^Marginal leaf cells differentiated, longer and narrower than within; leaves pointed atapex, margins recurved. ^  2392.^Leaves widest above the middle, not decurrent; costa in upper leaves excurrent in anawn. ^  B. capillare2.^Leaves widest below the middle, decurrent; costa ending before apex to shortlyexcurrent. ^  B. pseudotriquetrumCampylium1.^Costa absent, or short, usually ending below mid-leaf; leaves squarrose or wide-spreading, tapering from a cordate base into a long, fine point. ^ C. stellatum1. Costa extending to mid-leaf or beyond. ^  22. Leaves lanceolate from a broadly ovate base, lanceolate apex recurved.^  C. chrysophyllum2.^Leaves ovate-lanceolate, gradually tapering, straight and spreading. . .. C. polygamumClaopodium1.^Leaf cells multipapillose with two or more papillae per cell. ^ C. bolanderi1.^Leaf cells unipapillose ^  C. crispifoliumDicranella1.^Leaves lanceolate from a broad sheathing leaf base; leaf margins plane. D. schreberiana1.^Leaves lanceolate, gradually narrowed from base to apex, leaf base not sheathing; leafmargins recurved in lower leaf. ^  D. variaDicranum1.^Leaf cells papillose on the dorsal surface with one papilla per cell. . . D. fuscescens1. Leaf cells smooth ^  22. Leaf cells strongly pitted; leaves falcate-secund to straight and erect; leaf tips notusually broken; costa ending before apex to shortly excurrent. ^ D. scoparium2.^Leaf cells not pitted; leaves straight and erect; leaf tips often broken; costa longexcurrent. ^  D. tauricum40Didymodon1.^Leaf cells smooth. ^  D. rigidulus var. gracilis1. Leaf cells papillose  22. Ventral cells of the costa elongate at mid-leaf; leaves squarrose when moist.^  D. fallax var. reflexus2.^Ventral cells of the costa quadrate at mid -leaf; leaves spreading when moist.D^ vinealis var. flaccidusEncalypta1.^Plants with abundant gemmae clustered in leaf axils, gemmae consisting of brancheduniseriate filaments. ^  E. procera1. Plants without gemmae.  22. Leaf margins recurved at mid-leaf; dorsal surface of the costa smooth, or papilloseabove; capsule smooth, calyptra fringed, seta green for most of its length, red justbelow the capsule, peristome single, consisting of short red teeth. ^ E. ciliata2.^Leaf margins plane at mid-leaf; dorsal surface of the costa papillose by projecting ofcell ends; capsule ribbed, calyptra fringed or not. ^  33.^Capsule ribbed longitudinally; calyptra not fringed; seta red. ^ E. rhaptocarpa3.^Capsule (not observed) ribbed spirally; calyptra fringed. ^ E. proceraEurhynchium1.^Stem and branch leaves different in size but similar in shape; leaf apices more or lessacute to commonly obtuse or rounded (best seen in branch leaves). ^ 21.^Stem and branch leaves different in shape; leaf apices acute, never obtuse orrounded. ^  3In Eurhynchium praelongum and E. oreganum, stem and branch leaves are differentiated; branch leaves areovate, not markedly decurrent and not squarrose, while stem leaves are cordate with a very broad, more or lessauriculate base and a lanceolate, more or less squarrose apex, and are strongly decurrent. This difference in leafshape can be used to distinguish between branches and lateral stems.412.^Branch leaves usually < 1 mm ^ E. pulchellum var. pulchellum2. Branch leaves usually > 1 mm  E. pulchellum var. barnesii3. Plant usually slender; branching variable, regular or irregular; lateral stems usuallycommon on main stem; stems usually more or less sparsely and irregularly pinnate, ormore or less regularly and/or closely branched in which case commonly bipinnate;leaves not (or rarely slightly) plicate. ^  E. praelongum3.^Plant usually coarse; branching more or less regular; lateral stems usually few on mainstem; stem evenly and closely pinnate (mostly once pinnate, rarely sparsely bipinnate),somewhat frond-like; leaves, mostly those of stem and larger branches, often plicate.^  E. oreganumFissidens1.^Plant very small, less than 1 cm long; leaf margins bordered by elongate cells; plantsof dry habitats ^  F. bryoides1.^Plant usually more than 1 cm long; leaf margins not differentiated from the rest of thelamina; plants of wet habitats. ^  F. grandifronsHomalothecium1.^Leaf margins serrulate at apex, and occasionally also slightly serrulate at base. H fulgescens1.^Leaf margins toothed at apex and base, the teeth curved toward leaf base (hook-like).H nuttalliiHymenostylium1.^Leaves gradually lanceolate, curved toward the stem when dry, usually broadlyrecurved with the recurved portion several (2 or more) cells wide; costa usually 80 ormore gm wide at base; stem smooth. ^  H. insigne1.^Leaves linear, tapering into a point near apex, appressed to contorted when dry,narrowly recurved on one or both sides with the recurved portion 1 or 2 (occasionally3) cells wide; costa usually 40 to 60 gm wide at base; stem smooth or papillose.^  H recurvirostre42Hypnum1.^Cortical cells of stem thick-walled, similar to the cells within; leaves rounded atinsertion. ^  H. circinale1. Cortical cells of stem thin-walled, hyaline, larger than the cells within; leaves straightat insertion. ^  22. Alar cells differentiated, forming a distinct group of swollen cells. . . . . H. lindbergii2.^Mar cells not clearly differentiated, few, quadrate, or occasionally with a few,enlarged, hyaline cells at the alar margin. ^ H. subimponensMnium1.^Leaf margins with inner core of stereids visible in cross-section; leaf cells angular,walls not thickened at the corners; leaves not contorted when dry. . . M. spinulosum1. Leaf margins without an inner core of stereids; leaf cells rounded or somewhatangular, walls thickened or not at the corners; leaves contorted when dry. ^ 22. Median leaf cells mostly more than 20 pm (often up to 40), rounded, thickened at thecorners with the walls otherwise not markedly thickened; dioicous or synoicous. . . 32.^Median leaf cells mostly less than 20 pm (up to 25), rounded to somewhat angular,thickened or not at the corners with the walls generally thick and firm; dioicous. M. thomsonii3.^Synoicous; costa not toothed at the back near apex (look at several leaves). M. marginatum3.^Dioicous; costa usually toothed at back near apex. ^ M. ambiguumNeckera1.^Leaves abruptly rounded below the pointed apex; leaf apices serrulate; plant shiny;capsules immersed ^  N. pennata1.^Leaves only slightly rounded below the pointed apex; leaf apices toothed, the teethlong, sharp, and often recurved toward leaf base; plant dull; capsules exserted.N douglasii43Orthotrichum1.^Dioicous. ^  0. lyellii1. Autoicous, or sexuality unknown. ^  22. Leaves flexuose to crisped and contorted when dry; capsule exserted, with immersedstomata; endostome of 16 fine, linear (or hair-like) teeth; calyptra glabrous or hairy ^^  32.^Leaves erect, and appressed when dry; capsule immersed to shortly exserted; withsuperficial stomata; endostome of 8 or 16 lanceolate teeth; calyptra hairy. ^ 43.^Leaves flexuose when dry; stomata abundant, found on the middle and lower portionof the capsule; exostome of 8 whitish teeth (often split to 16 when dry). 0. consimile3. Leaves crisped and contorted when dry; stomata found on the middle and upperportion of the capsule; exostome of 16 reddish teeth. ^ 0. pulchellum4. Exostome and endostome of 8 teeth each; capsule emergent to shortly exserted,slightly ribbed below the mouth when dry. ^  0. speciosum4.^Exostome and endostome of 16 teeth each; capsule immersed to emergent, ribbed orsmooth when dry. ^  55.^Capsule ribbed when dry; dioicous. ^  0. lyellii5.^Capsule smooth when dry; autoicous; endostome teeth irregularly notched, ragged. 0. striatumPlagiomnium1.^Leaf margins toothed or entire, the teeth short and blunt; leaf apices rounded, usuallyapiculate; prostrate stems creeping on the substratum with rhizoids scatteredthroughout the length of the stem; leaves not or only slightly decurrent; operculumrostrate ^  P. rostratum1. Leaf margins toothed, the teeth sharp; leaf apices more or less pointed, apiculate;prostrate stems if present, usually arching with rhizoids at both ends; operculum conic.^  22. Plants with prostrate stems; leaves strongly decurrent; costa with stereids. . P. insigne2.^Plants without prostrate stems; leaves not or only slightly decurrent; costa withoutstereids. ^  P. venustum44Plagiothecium1.^Leaves mostly symmetric^ 21. Leaves asymmetric.  32. Leaves undulate, complanate; plants robust, whitish green. P. undulatum2. Leaves smooth, usually concave and not complanate (occasionally complanate).P cavtfolium3. Decuffent portion of leaves tapering, with the cells usually elongate, never swollen;median leaf cells narrow, less than 12 gm wide. ^ P. laetum3.^Decurrent portion of leaves often bulging, with the cells rounded, often swollen;median leaf cells wide, more than 12 gm wide. ^ P. denticulatumPohlia1.^Leaves with a metallic shine; leaf cells narrowly elongate, commonly over 10:1. P. cruda1. Leaves not shiny; leaf cells broadly elongate, less than 10:1 (except for marginalcells). ^  22. Leaf cells thick-walled; plant green to brownish; leaves erect, margins plane orrecurved, costa strong. ^  P. nutans2.^Leaf cells thin-walled, or not conspicuously thickened; plant whitish green; leafmargins plane. ^  P. wahlenbergiiPolytrichum1.^Leaf margins sharply toothed, and plane when moist; apical cells of the lamellaerounded in cross-section, with the walls more or less evenly thickened. . P. formosum1. Leaf margins entire, and incurved when moist; apical cells of the lamellae pear-shapedin cross-section, with the upper wall distinctly thickened. ^ 22. Costa long excurrent as a hyaline, toothed awn. ^ P. pihferum2.^Costa shortly excurrent as a reddish awn.  P. juniperinum45Porotrichum1.^Mar cells not differentiated from surrounding cells, elongate; leaf apices acute; leafmargins toothed in upper leaf with coarse sharp, irregular teeth. ^ P. bigelovii1.^Mar cells differentiated, quadrate; leaf apices acute to obtuse; leaf margins serrate toserrulate in upper leaf. ^  P. vancouverienseRacomitrium1.^Leaves with hyaline hair-points. ^  21. Leaves without hair-points.  72. Hair-points papillose ^  32.^Hair-points usually denticulate, but not papillose. ^  43.^Leaf cells smooth (with the exception of the hair-point). ^ R. lanuginosum3. Leaf cells papillose ^  44. Costa ending around mid-leaf; marginal leaf cells above the alar cells usually elongateand thin-walled. ^  R. canescens4.^Costa ending near apex; marginal leaf cells above the alar cells short (usuallyquadrate) and thick-walled. ^  R. elongatum5.^Leaves bordered at the basal margin by elongate, esinuose, thin-walled, more or lesshyaline cells, and alar cells differentiated, smooth, esinuose and thin-walled; leaf cellspapillose, with several low papillae per cell. ^  R varium5. Leaves not bordered at the basal margins and alar cells not differentiated, or if so, leafcells smooth ^  66. Hair-points strongly decurrent, not strongly denticulate; alar and supra-alar marginalcells not differentiated; leaves linear-lanceolate; costa usually bistratose, stronglycanaliculate; leaf cross-sections at mid-leaf V-shaped and showing round, thick-walledlaminal cells; upper marginal leaf cells elongate ^ R. lawtonae6.^Hair-points shortly decurrent or abruptly ending , denticulate to spinulose; alar andsupra-alar marginal cells differentiated or not; leaves linear-lanceolate to ovate-lanceolate; costa usually 2 to 4-stratose, strongly canaliculate or not; leaf cross-sections at mid-leaf variable, and showing round or elliptic laminal cells; uppermarginal leaf cells short to shortly elongate   R. heterosti chum467.^Apical leaf cells elongate (more than 3:1), apex narrowly long lanceolate; in section,papillae seen as large rounded thickenings on the cross-walls. ^ R. fasciculare7. Apical leaf cells short (less than 3:1) or mixed. ^  88. Leaves ungulate, broadly obtuse to rounded at apex; leaf margins usually with bluntteeth toward apex ^  R. aciculare8.^Leaves ovate-lanceolate, narrowly obtuse to acute at apex; leaf margins not toothed,smooth or finely crenulate at apex. ^  99.^Leaf cells with several low papillae per cell. ^  R. varium9.^Leaf cells smooth. ^  R. pacificumRhytidiadeOhus1.^Leaf cells in the upper portion of the leaves papillose on the back by projecting cellends; costa long and double, usually extending beyond mid-leaf. ^ R. triquetrus1. Leaf cells smooth throughout; costa absent, or short and double, ending below mid-leaf. ^  22. Leaves plicate; alar cells not differentiated. ^  R. loreus2.^Leaves not plicate; alar cells differentiated, shorter, wider, and hyaline.R squarrosusScleropodium1.^Basal leaf cells (last few rows) elongate; median leaf cells very long, often to 15-20:1. S. touretii var. colpophyllum1.^Basal leaf cells (last few rows) short, rounded, quadrate or short rectangular; medianleaf cells shorter, usually around 10:1.^ S. cespitansSeligeria1.^Peristome absent. S. donniana1.^Peristome present ^  2472.^Capsule hemispherical; seta straight when moist; columella long, often retaining theoperculum once detached or sticking out after the latter has fallen off; spores large(18-32 gm). ^  S. tristichoides2.^Capsule ovoid; seta usually curved when moist; columella short; spores small (8-18Pm).^ S campylopodaSphagnum1.^Plants green, never with red coloration; stem leaves ungulate, broadly truncate, apexfringed. ^  S. girgensohnii1.^Plants often red coloured; stem leaves triangular, apex narrow, often involute.S capilhfoliumTetraphis1.^Seta straight and smooth ^  T. pellucida1.^Seta bent in the middle, and papillose above the bend. ^ T. geniculataTortula1.^Costa excurrent as a coloured point or short awn, not readily visible at lowmagnification (10x); leaf margins differentiated at mid-leaf, cells longer and narrowerthan within, smooth or with low papillae. ^  T. subulata1. Costa usually long excuffent as a white (often reddish at the base), denticulate awnreadily visible at low magnification (10x); leaf margins not differentiated. ^ 22. Leaves erect-spreading when moist, more or less straight, not curved. . . T. princeps2.^Leaves recurved when moist, curved backward, apices pointed down toward the baseof the plant. ^  T. ruralisUlota1.^Spores large, more than 30 pm in diameter. ^ U. megalospora1.^Spores small, less than 30 pm in diameter.  U. obtusioscula482.2.3. Notes on problematic taxaAtrichumAtrichum selwynii and A. undulatum are both common species in southwestern BritishColumbia (Lawton 1971, Vitt et al. 1988). They are thus both likely to be found at BridalVeil Falls. The main difference between the two species involves their sexuality. A. selwyniiis always dioicous, while A. undulatum can be either monoicous or dioicous (Ireland 1969a).Treatments of the genus by Ireland (1969a, 1971) suggested a series of other characters todifferentiate the two species. Such characters involve the nature of the calyptra, the curvatureof the capsule, the number of sporophytes borne on each plant, and the characteristics of theleaf cells. However, based on observation of my material and of herbarium specimens, thesecharacters appear variable and greatly overlapping. Hispid-tipped calyptrae, which Irelandattributed to A. undulatum, were found to occur in dioicous material of A. selwynii. Curvatureof the capsule was variable even within a single population. Similarly, numbers ofsporophytes per plant varied between populations of the same species. Finally, Irelandattributed smaller, thick-walled, collenchymatous, rounded leaf cells to A. undulatum, andlarger, thin-walled, non-collenchymatous, angular leaf cells to A. selwynii. Thischaracterization, though observable, was not consistent in our material. The nature of the leafcells often varied between leaves of the same plant and even within a single leaf.Monoicous sexuality remained the only character that positively distinguished A.undulatum from A. selwynii. In the case of dioicous specimens or of those that could not besexed, careful observation of the nature of the leaf cells appeared to be the preferablesolution.49Only one specimen collected during the course of this study was monoicous,confirming the presence of A. undulatum at Bridal Veil Falls. Other specimens were eitherseemingly dioicous, or did not bear sporophytes and/or sex organs at the appropriate stage forassessing sexuality. They were identified based on the characteristics of leaf cells, asmentioned above.Dieranella schreberiana versus Trichodon cylindricusSterile material of Dicranella schreberiana is not easily distinguishable fromTrichodon cylindricus. Furthermore, the two species occur in the same types of habitat. Dixonand Jameson (1924), Smith (1978), and Crum and Anderson (1981) suggested the roughnessof the subula as a discriminant character. The subula is said to be toothed-papillose all aroundin Trichodon cylindricus, while in Dicranella schreberiana, the teeth should be found only onthe margin of the subula. Lawton (1971) also indicated a difference in size between the twospecies. According to her descriptions, stems of Dicranella schreberiana reach up to 30 mmin height, while Trichodon cylindricus is smaller, rarely reaching more than 5mm in height.The presence of both species at Bridal Veil Falls was confirmed by sporophyticmaterial of Trichodon cylindricus, and by material with the long shoots attributable toDicranella schreberiana. As a guide for the identification of small gametophytic material alsocollected, we examined and compared the roughness of the subula between the latterspecimens. We were unable, however, to observe the difference in roughness described above.50All sterile material collected during the course of this study was therefore classified asDicranella schreberiana.PlagiochiktGodfrey (1977) derived a key to separate Plagiochila asplenioides from P. satoi frompapers by Inoue and Schuster. The characters use in that key proved to be greatly variable.Although this key worked well for some specimens, Godfrey encountered many transitionalspecimens. She concluded that there were no satisfactory means of separating the two speciesin southwestern British Columbia. Specimens belonging to this problematic group weretreated as the P. asplenioides complex.Godfrey (1977) pointed out that a detailed study of the group was needed to solve theproblem. Hong (1992), who worked on the genus Plagiochila in Western North America,proposed key characters which are essentially the same as those unsuccessfully used byGodfrey. The problem thus remains unsolved.To avoid problems with transitional specimens, Godfrey's treatment was adopted here:species of the P. asplenioides/P.satoi group were identified as P. asplenioides sensu lato.PlagiomniumTaxonomic treatments of Plagiomnium (Lawton 1971; Koponen 1974) are basedprimarily on fertile material. They refer to characteristics of leaves of mature material,sexuality, and sporophytic characters. Neither keys nor descriptions account for differentgrowth forms, life stages or environmental variants. In the course of this study, we51encountered difficulties identifying sterile and plagiotropous material.The ecological aspects of the study, required the collection of material to record itspresence in a certain habitat, rather than the collection of ideal material for the purpose oftaxonomic work. In general, taxonomic treatments that treat variability would undoubtedly beuseful in ecological work.RacomitriumThe classification of Racomitrium sect. Laevifolia (or heterostichum group) isproblematic, reflecting the inherent variability of this taxon. A complete and thoroughtaxonomic treatment of this complex had been lacking. Recently, the taxonomy of the groupwas revised by Frisvoll (1988). His treatment has been used here as a reference for theidentification of the specimens of the heterosti chum group found at Bridal Veil Falls.Frisvoll (1988) based his treatment on taxonomic characteristics which he recognizedas stable and therefore useful for species recognition. They include the structure of the costa,the alar cells and supra-alar marginal cells, the leaf margin, and the perichaetial leaves. Basedupon Frisvoll's treatment we were able to recognize two species of the heterostichum groupin the Bridal Veil Falls collection: Racomitrium pacificum, and R. lawtonae. The former ischaracterized by the absence of hair-points and unistratose leaf margins. The latter ischaracterized by its long lanceolate leaves with wealdy denticulate, long decurrent hair-points,unistratose margins, undifferentiated alar cells, elongate upper marginal cells, median leafcells rounded in cross-section, and bistratose, canaliculate costae (Ireland 1970, 1976; Frisvoll1988).52My material also included several specimens which did not conform satisfactorily tothe description of either of the above species. We encountered the same difficulties of highvariability mentioned by other taxonomists (Frisvoll 1988). Unfortunately, I had at mydisposal only very few specimens identified by Frisvoll. Such specimens would have helpedclarify the concepts developed by Frisvoll (1988) and familiarize ourselves with the variabilitypermitted within the different species concepts.The specimens that would not fit into the segregate species were classified asRacomitrium heterostichum sensu lato. They usually possessed strongly denticulate hair-pointsthat were not or only shortly decurrent, short to slightly elongated upper marginal leaf cells,median leaf cells elliptic in cross-section, and predominantly unistratose leaf margins. Thespecimens probably include some material of Racomitrium heterostichum (e.g. Djan-Chëlcar &Dagenais 91-106) with a costa broad and bistratose at mid-leaf, and of R. affine (e.g. Djan-Chëkar & Schofield 91-544; Djan-Chdlcar & Reid 91-69) with a costa narrower and tri-stratose at mid-leaf. Our concept of these taxa, however, is not sufficiently clear to positivelyidentify them as such. Some specimens might also be variations of Racomitrium lawtonae(e.g. UBC b20413, b20439, and b20418). These specimens typically possessed bistratose,canaliculate costae, unistratose margins and undifferentiated alar cells, but hair-points wererelatively short decunent, upper marginal leaf cells were short, or some differentiated cellsoccurred at the basal margin. Among all specimens encountered unistratose leaf marginspredominated. This eliminates as possible species, Racomitrium brevipes, R. occidentale, andR. obesum.532.2.4. Barbula convoluta var. gallinula, new to British Columbia.Barbula convoluta var. gallinula Zander is reported here as new to British Columbia.Three specimens were collected during the course of this study and a duplicate of one ofthem was sent to R.H. Zander for confirmation.While Barbula convoluta var. convoluta is common and widely distributed (Smith1978, Zander 1979), var. gallinula has been reported only from the Northwest Territories,Yukon and Alaska (Zander 1979, and pers. communication (1992)). Southwestern BritishColumbia becomes the southernmost known locality for this variety. The wide gap in itsdistribution is most likely to be the result of a lack of collection of this small andinconspicuous plant.54Chapter 3. Ecology3.1. Introduction - Species diversityThe species diversity of a community is normally expressed in terms of number ofspecies present (species richness) and in terms of the relative abundance of these species(evenness). The results of the present study include estimates of species richness; relativeabundance was not measured with any precision. Therefore, any reference to diversity alludesto species richness only. Species richness by itself can be considered as a simple, unweighedmeasure of diversity (Pielou 1975).Several hypotheses have been proposed to indicate which community factors determinediversity. Such factors include: time, productivity, succession, stability, environmentalextremes, predictability, competition, and habitat complexity and heterogeneity (Slack 1977;Putman and Wratten 1984). As noted by Pielou (1975), some factors play a role at a localscale, that is in a particular community at a particular time. These small scale, short termfactors are the proximate causes of diversity. They are the concern of ecology. Other factorsrelated to diversity play a role over areas of wider geographic extent and over considerableperiods of time. These large scale, long term factors are the ultimate causes of diversity. Theyare the concern of biogeography and are considered in Chapter 4.More specifically, this chapter emphasizes the role of habitat complexity andheterogeneity as a promoter of diversity. Substratum-patchiness, for instance, has beenassociated with increased diversity of small organisms, such as bryophytes (Pielou 1975).Because of their small size, bryophytes are affected by very local variations in environmentalconditions.553.2. Materials and methodsThe data used for this general analysis of bryophyte ecology at Bridal Veil Falls werederived from ecological information recorded when specimens were collected (see Section2.1.1). Therefore, only specimens collected during the course of this study were included inthe analysis. As specimens were determined, other bryophytes found in mixture with theprincipal species were also identified and recorded. A total of 807 records of mosses and 189records of liverworts were compiled for this ecological analysis. Eighty-three percent of thebryophytes composing the flora are represented. They are listed in Appendix C.3.2.1. Definition of habitat categoriesFor the purpose of this analysis, habitats where bryophytes were collected wereclassified into six different categories. These different categories were defined as follows (seeFigure 1.2):1) Anthropogenic habitats: Anthropogenic habitats are the direct result of humanactivity. They comprise the park installations, parking lot, trails, lawn and picnic area, and aclear-cut strip, or meadow, over a buried pipeline.2) Forest habitats: Most forested sites found throughout the study area are treated asforest habitats. Also included within the forest habitats are edges of the forest and the forestedscree slopes found at the base of the cliffs. The exceptions are the forested cliffs, streams,spray-zones and edges of the boulder slope which are treated as other habitat categories.3) Boulder slope habitats: Habitats within this category are all part of the boulderslope west of the park. This includes not only the open portion of the slope, but also theforested edges and a shrubby area at the top.564) Cliff habitats: Cliff habitats include the series of cliffs that define the southernborder of the study area and a few vertical outcrops found in the forest below. They compriseopen cliffs and also those shaded by the adjacent forest, as well as the sites of waterfalls.5) Stream habitats: Stream habitats encompass the banks and beds of the four foreststreams found in the study area. This includes an intermittent stream which dries up duringthe summer months.6) Spray-zone habitats: This category includes areas at the base of waterfalls whichreceive spray from the falls throughout most of the year (during the summer months, waterlevels being low, spray can be much reduced). Excluded from this category are cliffs withinthe spray zones; these are classified as cliff habitats.3.2.2. Definition of substratum categoriesWithin each type of habitat, bryophytes occur on a variety of substrata. The differentsubstrata were further classified independent of habitat. Substratum categories were:1) Rock: Rock as a substratum comprises cliffs, boulders, smaller rocks, projectingoutcrops and gravel. It can include vertical surfaces such as cliffs and boulder faces,horizontal surfaces such as rocks and projecting outcrops on the forest floor or the crests ofboulders, or finally loose surfaces such as gravel. Rocks found in the study area were dividedinto two types: calcareous and non-calcareous.2) Vegetation: Trees and shrubs in the study area support epiphytic communities ofbryophytes. Some bryophytes were also found to grow on other species of bryophytes.Epiphytes were classified on the basis of the species on which they were collected, as well as57on the basis of where they grew on the tree or shrub. The roots and the base of the tree (>25cm from the ground) were considered as one category, and the trunk another. Branches in theunderstory, as well as branches and stems of shrubs, formed a third category. Branches of thecanopy, accessible from recently-fallen trees and branches, formed the last category.3) Soil: Soil as a substratum consists of either the upper surface of mineral soil, soilaccumulated over rock, soil accumulated between rocks, or earth banks. In any of these cases,the soil involved can be of different types. Those encountered in this study were classified aslitter, organic (which includes humus), sand, clay, and silt.4) Wood: Wood in various stages of decay is also an important substratum forbryophytes. It is represented by logs, stumps or snags, and branches. Three different states ofdecay were recognized in this study: recently-fallen (still with bark), decaying (has lost itsbark, but is still solid), and finally rotten (falls apart easily).3.2.3. Similarity indicesSimilarity indices were used to determine the degree of similarity between pairs ofhabitats in terms of the species composition of their respective floras. The Jaccard Index ofSimilarity, which is based on binary (or presence/absence) data, was chosen for this purpose.This index is the proportion of species which co-occur in two sites to the total number ofspecies found in at least one of the two sites, or:JI = aa+b+cwhere a is the number of species occurring at both sites, b is the number of species occurringonly in the first site, and c is the number of species occurring only at the second site (Digby58and Kempton 1987; Ludwig and Reynolds 1988). The minimum value for this index is 0, thatis, when the two sites have no species in common. The maximum value is 1, that is, when thetwo sites have identical floras. Conjoint absences of species are ignored in this index.The relationship between habitats based on similarity in species composition wasrepresented graphically using multidimensional scaling (Wilkinson 1990).3.3. Results and discussion3.3.1. Floristic diversity in terms of habitats and substrataThe habitats recognized at Bridal Veil Falls can be separated into two groups based onthe number of bryophyte species they contain (Table 3.1): boulder slope, cliff and foresthabitats are more diverse, whereas, anthropogenic, stream and spray zone habitats are lessdiverse. Such results reflect, at least in part, the relative area occupied by each of thesehabitats at Bridal Veil Falls. Although precise measurements are not available, the forestcertainly occupies the largest area. Cliffs and boulder slope are also relatively extensive.Anthropogenic habitats are probably comparable in size to the latter. On the other hand,streams and spray zones both occupy much smaller areas than any of the other habitats.Furthermore, sampling was focused on the largest habitats, those where bryophytes were mostabundant, and those that seemed, at first glance, the most diverse - in other words, forest,cliffs and boulder slope. However, area and sampling effort are not the only factors affectingdiversity. The following discussion will outline how other factors also affect the number ofspecies observed in the different habitats.59Table 3.1. The distribution of bryophytes between habitats, between substrata, and betweensubstrata within each habitat. (Total numbers of bryophytes are given in bold, numbers ofmosses are in parentheses 0, numbers of liverworts are in brackets H.)SubstrataVegetation Rock Soil WoodHabitat Anthropogenic 0 7 15 0 20types (0) (7) (15) (0) (20)[0] [0] [0] [0] [0]Boulder slope 1 54 30 13 78(1) (46) (21) (8) (61)[0] [8] [8] [8] [17]Cliff 1 61 43 0 78(1) (48) (30) (0) (60)[0] [13] [13] [0] [18]Forest 60 14 24 45 88(46) (14) (23) (32) (67)[14] [0] [1] [13] [21]Stream 5 18 12 18 35(5) (16) (8) (16) (29)[0] [2] [4] [2] [6]Spray zone 3 9 8 21 33(3) (8) (7) (13) (25)[0] [1] [1] [8] [8]64 107 94 73 Total(50) (88) (73) (52) number[14] [19] [21] [21] ofspecies''Species totals are independent of the body of the table. They report total species number on each habitat orsubstratum. As species can occur on more than one habitat/substratum combination, column or row totals willalways be less than or equal to species totals.60In terms of substrata, the highest species diversity was found on rocks and on soil,while diversity on vegetation and on wood is lower (Table 3.1). Such results are notsurprising considering that rock and soil are the most important substrata both in the boulderslope and on cliffs, two of the three most diverse habitats (Table 3.1). Combined, the flora ofthe boulder slope and cliffs comprise 71% of all species found at Bridal Veil Falls.It should be noted that 82% of the species growing on soil throughout the study areaoccur on soil over rock. Soil in cliff and boulder slope habitats, particularly, is usually soilover rock. Rock thus serves as a secondary substratum. This further emphasizes theimportance of rock as a substratum at Bridal Veil Falls.The differences observed between substrata in terms of species diversity are also partlya reflection of the general ecology of bryophytes. For instance, Smith (1982) observed thatthe number of epiliths (species growing on rock) in a flora is usually greater than the numberof epiphytes - this applies to both obligate or facultative epiliths or epiphytes. Corroboratingthis observation, Schofield (1976) reported that the majority of moss genera in BritishColumbia are able to grow on rock.3.3.2. Widely distributed versus restricted speciesAmong the bryophytes found at Bridal Veil Falls, some species occurred in a widevariety of habitats and substrata, while others were restricted to a certain type of habitat orsubstratum. Examples of widely distributed taxa are: among mosses, Claopodium crispifolium,Eurhynchium praelon gum, Homalothecium fulgescens, Isothecium myosuroides, Mniummarginatum, M. thomsonii, Plagiomnium rostratum, Rhytidiadelphus triquetrus, and Tortella61tortuosa, and among liverworts, Conocephalum conicum, Plagiochila asplenioides s.L, andScapania americana. Examples of species restricted (or unique) to a certain substratum in acertain habitat are: for the mosses, Seligeria spp., Hypnum lindbergii, Racomitrium elongatum,R. canescens, Bryum miniatum, Buxbaumia piperi, Dicranella varia, Platyhypnidiumriparioides, and Ulota spp., and for the liverworts, Preissia quadrata, PoreIla cordaeana,Anastrophyllum minutum, Gymnomitrion obtusum, Moerckia hibernica, and Lepidozia reptans.Species occurring in a wide variety of habitats and substrata can be regarded asspecies with a wide range of ecological tolerance, that is, in terms of the differentmicroenvironments found at Bridal Veil Falls. On the other hand, species unique to a certainhabitat or substratum can be regarded as species with a relatively limited range of ecologicaltolerance (again in terms of the different microenvironments found at Bridal Veil Falls only).Evidently, there exists a gradient of ecological tolerance among species. The majority ofspecies have varying degrees of ecological tolerance intermediate to that of the speciesmentioned above.The restriction of a species to a certain type of habitat or substratum is considered as akey element of this study. The exclusive presence of a species in a certain habitat confers awell defined role to this particular habitat as a determinant of diversity in the study area. Italso provides an indication of how this habitat might differ from the others. Among thespecies found at Bridal Veil Falls, a high proportion were restricted to a certain habitat or to acertain substratum - respectively 45% and 43% (Table 3.2 and 3.3). This illustrates howvariety in terms of habitat is important to the diversity of an area.62Table 3.2. Number of bryophytes unique to each habitat.Habitat types Number of bryophytes unique to each habitat % oftheentireflora ofeachhabitatMosses Liverworts TotalAnthropogenic 6 0 6 30Boulder slope 15 8 23 29Cliff 10 4 14 18Forest 21 6 27 31Stream 4 1 5 14Spray zone 2 2 4 12Total 58 21 79 4563Table 3.3. Number of bryophytes unique to each substratum.Substrata Number of bryophytes unique to each substratum % of theentireflora ofeachsubstrataMosses Liverworts TotalVegetation 5 3 8 12.5Rock 26 7 33 31Soil 17 4 21 22Wood 7 7 14 19Total 55 21 76 4364The number of species restricted to each habitat (Table 3.2) reflects species diversitywithin each habitat (Table 3.1). The most diverse habitats (forest, boulder slope and cliff) hadthe largest number of unique species, while few species were restricted to habitats with lowspecies diversity (anthropogenic, stream and spray zone). However, closer examination of theresults shows that high species diversity and large number of unique species are notnecessarily correlated. This is best illustrated by the percentage of species in the flora of eachhabitat, confined to that habitat (Table 3.2). Both forest and boulder slope, which had highspecies diversity, had a relatively high proportion of unique species, and both stream andspray zone, which had low species diversity, had a relatively low proportion of uniquespecies. On the other hand, cliff habitats, despite high species diversity, had a relatively lowproportion of unique species, and anthropogenic habitats, the least diverse, had a relativelyhigh proportion of unique species.The number of species unique to each substratum (Table 3.3) also reflects speciesdiversity (Table 3.1). In this case however, the pattern is consistent. Substrata with highspecies diversity (rock and soil) had a large number of unique species, while few species wererestricted to the less diverse substrata (vegetation and wood). The substratum with the highestpercentage of unique species was rock. Vegetation had the lowest percentage. Again, theseresults corroborate observations by Smith (1982) who noted that the number of obligateepiliths in a flora is usually larger than the number of obligate epiphytes. Schofield (1976)made similar observations, noting that few mosses in British Columbia's flora are confined toan epiphytic habitat.653.3.3. Floristic similarity between habitatsTable 3.4 describes the similarity in species composition between the moss flora ofeach pair of habitats using the Jaccard Index of Similarity. None of the indices of similarityobtained are particularly high, ranging only between 0.09 and 0.31. This indicates that thehabitat categories defined for this project generally discriminate relatively well between actualhabitats occupied by bryophytes. This is not to say that overlap does not occur, or that it hasbeen reduced to a minimum. To do so a much larger number of categories, as well as toolssuch as statistical classification of communities, would be required. Classification would alsoneed to be done at a much smaller scale. The categories used here are artificial, definedmainly for the purpose of a broad discussion of ecology.Among the indices of similarity obtained, there are nevertheless some relativelyimportant differences. This is best illustrated in Figure 3.1 which provides a graphicrepresentation of the similarity matrix given in Table 3.4.Anthropogenic habitats had among the lowest indices of similarity for all habitats(Table 3.4). In other words, anthropogenic habitats shared only a small portion of their florawith each of the others. Such results reflect the fact that anthropogenic habitats have thelowest diversity of mosses (Table 3.1), absence of liverworts, as well as one of the highestpercentage of unique species (Table 3.2).Cliffs, forest and boulder slope show a relatively high degree of similarity. This is notsurprising considering that these three habitats have the highest species diversity (Table 3.1).Cliff, forest and boulder slope habitats all have a large number of unique species (Table 3.2).Nevertheless, the results indicate that a large number of widely occurring species would also66Table 3.4. Similarity between habitats based on composition of the moss flora using theJaccard Index of Similarity.Habitat types BoulderslopeCliff Forest Stream SprayzoneAnthropogenic 0.09 0.14 0.13 0.17 0.10Boulder slope 0.27 0.31 0.14 0.10Cliff 0.22 0.24 0.25Forest 0.17 0.12Stream 0.2367Figure 3.1. Relationship between habitats based on similarity in species composition usingmultidimensional scaling (stress value of 0.000281).21Dimension 2 0-1-2^1 ^ i^ 1_o AnthropogenicForest o^_^Streams o^ o Boulder slopeo Cliffs^-^Spray zones o_I-2^ -1^ 0^ 1^ 2Dimension 1'Stress is very low indicating that this graphic representation fits the data generally well. The Shepard diagram(not shown), however, resembles a stepwise function indicating that a degenerate solution was achieved. Therefore,although the general picture is appropriate, the graphical distances between points are not an entirely accuratedepiction of the Jaccard similarities between habitats. The use of a three-dimensional plot did not sensibly correctthis problem.68be shared between them. Examples would be Claopodium crispifolium, Homalotheciumfulgescens, Hylocomnium splendens, Hypnum subimponens, Isothecium myosuroides, Neckeradouglasii, Plagiomnium rostratum, Rhytidiadelphus triquetrus, Scleropodium cespitans,Conocephalum conicum, and Scapania americana, all of which occurred in all three habitats.Several others occurred in two of the three. Between boulder slope and cliff habitats, epilithiccommunities shared the largest proportion of species, with a coefficient of similarity of 0.25.Such a result is expected since rock is the predominant substratum in both of these habitats.Examples of species found on rock in both cliff and boulder slope habitats were Anacoliamenziesii, Bartramia pomiformis, Brachythecium plumosum, Grimmia torquata,Gymnostomum aeruginosum, Heterocladium macounii, Plagiopus oederiana, Polytrichastrumalpinum, Racomitrium heterostichum Si, Herbertus aduncus, and Marsupella emarginata.Between the boulder slope and the forest, the substrata sharing the largest number of specieswere rock in the boulder slope and forest vegetation, with a coefficient of similarity of 0.23.Here again both of these substrata are predominant in their respective habitat. Furthermore,there are many bryophyte genera in which species are able to grow either as epiphytes orepiliths (Schofield 1976). Examples of such species found both in the forest and on theboulder slope are: Antitrichia spp., Dicranum tauricum, Plagiomnium venustum,Pseudotaxiphyllum elegans, and Rhytidiadelphus loreus. Between cliff and forest habitats, nopairs of substrata showed a particularly high degree of similarity in species composition oftheir respective flora. The highest degree of similarity was a coefficient of 0.17 between rockin cliffs and forest vegetation. Again, this result points out how a portion of epiphytic speciesare often found growing on rock surfaces. Examples of species found in both cliff and forest69habitats are: Brachythecium asperrimum, Dicranella schreberiana, Eurhynchium pulchellumvar. pulchellum, Metaneckera menziesii, Platydictya jungermannioides, Thamnobryumneckeroides, Apometzgeria pubescens, Plagiochila asplenoides s.1., and Porella roellii.Stream, cliff and spray zone habitats also show a relatively high degree of similarity.This could be attributed to these habitats harbouring, at least in part, high-moisturemicroenvironments. Wide-ranging mesic and hydric species, such as Hygrohypnum luridum,Hymenostylium insigne, Fissidens grandifrons, Campylium stellatum, and Lophozia gillmanii,are more likely to be shared between streams, cliffs and spray zones than with any othertypes of habitat. This is illustrated by the low degree of similarity that streams and sprayzones have with forest and boulder slope. As described above, cliffs, nevertheless, have ahigher degree of similarity with forest and boulder slope, for they include a large portion ofdrier microenvironments as well as high-moisture ones. Combined with the high speciesdiversity observed on cliffs (Table 3.1), the above results outline the richness of cliffs interms of microhabitats. Furthermore, the relatively large proportion of species shared by cliffhabitats with each of the others, except anthropogenic ones, explains the relatively lowproportion of species to be found only on cliffs (Table 3.2).3.3.4. The occurrence of species in habitats and on substrataThe following examines the flora of the different habitats and substrata in more detailand provides further discussion of the results given above.70AnthroDogenic habitats The results have shown that the species composition of anthropogenic habitats isunique to those habitats. This is related to the uniqueness of anthropogenic habitats among allhabitats in the area in having soil - in this case bare ground - as a dominant substratum. Asshown in Table 3.1, soil was the important substratum in anthropogenic habitats in terms ofspecies diversity. Seventy-five percent of all the species found in this habitat grew on soil.Furthermore, nearly all the species unique to this habitat were growing on soil. They areAtrichum selwynii, Calliergonella cuspidata, Funaria hygrometrica, Hypnum lindbergii, andRhytidiadelphus squarrosus.The importance of soil as a substratum in anthropogenic habitats reflects an early stagein plant succession. Schofield (1985) pointed out that the persistence of bryophytes on soil issometimes brief. Successional changes usually permit colonization by seed plants whichoutcompete the bryophytes present in earlier stages. This is what is happening in the meadow(the area cleared for a pipeline). As weeds and grasses have grown more abundant at the site,the abundance and variety of bryophytes has declined (W.B. Schofield, pers. communication).On the other hand, other anthropogenic sites are kept at early successional stages bycontinued use and maintenance by humans.Sauer (1988) commented on the colonization of abandoned fields. His comments applyto the meadow and seem to apply also to other artificially modified anthropogenic habitats,such as the lawn and picnic area. He suggested that abandoned fields are invaded fromoutside rather than reclaimed by species in situ. His observations are for seed plants, but seemto apply to bryophytes as well. Considering dispersal distances, the most likely invaders of71anthropogenic habitats are species from the adjacent forest. However, it is likely thatappropriate microhabitats for forest species are scarce in anthropogenic habitats. Unlike theforest, the soil here does not include a thick layer of organic matter. The habitat being moreopen, light and moisture conditions also often differ, especially away from the edges betweenthe two habitats. Nevertheless, some of the more widely distributed species occurring in theforest are present also in anthropogenic habitats. They are Bryum capillare, Claopodiumcrispifolium, Eurhynchium praelongum, Homalothecium fulgescens, Plagiomnium insigne, P.rostratum, and Rhytidiadelphus triquetrus. On the other hand, we also observed a highproportion of weedy species in anthropogenic habitats, such as Atrichum selwynii,Calliergonella cuspidata, Funaria hygrometrica, and Rhytidiadelphus squarrosus. Amongthem are the species restricted to anthropogenic habitats. Presumably, they invaded fromlawns and fields or other disturbed habitats in the vicinity. It is possible that they wereaccidentally introduced from other lawns via moss fragments on lawnmowers.Boulder slope The boulder slope was found to be among the habitats with high species diversity, aswell as a high proportion of unique species. To some extent, such results can be related to thevery different environmental conditions found in the boulder slope in contrast to otherhabitats. Unlike any of the others, the boulder slope is open and exposed. Consequently, itexperiences much stronger extremes in terms of climatic conditions. Of the species occurringat Bridal Veil Falls exclusively in the boulder slope, several are species tolerant of longperiods of drought. Examples would be species like: Amphidium californicum, Pogonatum72umigerum, Polytri chum piliferum, Racomitrium spp., and Gymnomitrion obtusum.The boulder slope also differs edaphically from the other habitats. The rock is non-calcareous while in the rest of the study area, calcareous rock dominates. The only exceptionis a portion of cliff just above the slope where the rock surface is also non-calcareous.Therefore, several species restricted to the boulder slope and the cliff above are usuallyassociated with acidic substrata. Examples are: Encalypta ciliata, Pleurozium schreberi,Racomitrium spp., Sphagnum spp., Anastrophyllum spp., Frullania californica, Gymnomitrionobtusum, and Herbertus aduncus. Nevertheless, some species usually associated withcalcareous substrata also were found to occur in the boulder slope and the cliff above.Examples are species such as: Barbula convoluta var. gallinula, Encalypta rhaptocarpa,Gymrzostomum aeruginosum, Palustriella commutata, Plagiopus oederiana, Timmia austriaca,Tortella tortuosa, Tortula ruralis, Conocephalum conicum, and Preissia quadrata.It has been found that other elements may partially replace calcium (Brown 1982).Species usually associated with calcium could tolerate low calcium environments bysubstituting calcium with other elements. This might be the case for some of the speciesmentioned above. Among species where such a substitution has been observed, Brownreported Palustriella commutata. Another possible explanation is that run-off of calcium-richwater occurs over the rock where these bryophytes grew. It was mentioned previously that thewater in the area has a very high concentration of calcium. It is probable, especially on cliffs,that water running over the non-calcareous rock could provide the appropriate amounts ofcalcium for the growth of calcicoles. This argument is supported by the presence of speciespredominantly associated with calcareous rock on tree trunks or wood at the base of the cliffs73or in spray zones. Examples are Encalypta procera, Palustriella commutata, Timmiaaustriaca, and TorteIla tortuosa.Another important factor, with regard to high species diversity in the boulder slope, ismicrotopography. The large size of the boulders and their arrangement makes for verycomplex relief. It creates a wide variety of conditions in terms of exposure and consequently,a wide range of temperature and moisture conditions. Such differentiation in environmentalconditions also occurs between the open part of the slope and the edges. It is interesting tonote here that regardless of the substratum, most of the 36 species found in both the forestand the boulder slope were restricted in the boulder slope to partially or fully shaded sites,such as edges of the slope, north faces of boulders, overhangs, or depressions amongboulders. Examples are Rhytidiadelphus triquetrus, Pseudotaxiphyllum elegans, Hypnumsubimponens, TorteIla tortuosa, Bryum capillare, Jamesoniella autumnalis, Conocephalumconicum, and Scapania umbrosa. Exceptions, or species that were found in open microhabitatsin the boulder slope, are: the widely occurring Hylocomium splendens; Dicranoweisia cirratawhich occurred in the forest in the canopy of cedar where humidity is low and light intensityhigh in comparison to lower on the trees (Smith 1982); Polytri chum juniperinum found onsoil at the forest edge, again a presumably drier and more illuminated habitat; andAulocomnium androgynum, Scapania americana, and Lophozia ventricosa, all three occurringon wood in the forest.The assemblage of species found on wood in the boulder slope differed from that onwood in the other habitats: coefficients of similarity were 0.08 with wood in forest, 0 withwood in streams, and 0.05 with wood in spray zones. This could reflect the difference in74moisture and light conditions between the boulder and the three other habitats. During thesummer, temperatures remain cooler in the partial or full shade of the forest. Therefore, woodin forest or stream habitats retains some degree of moisture throughout most of the year. Thesituation is somewhat similar in spray zones which can be more open, but nevertheless arekept cool and moist by the spray. On the other hand, the open boulder slope becomes dryduring the summer months, as moisture quickly evaporates on warm and sunny days.However, only the liverwort Odontochisma denudatum was found to grow exclusively onwood in the boulder slope. The presence of this species in the boulder slope and at BridalVeil Falls will be discussed in the following chapter. Most of the other species that weregrowing on wood in the boulder slope were also found either as forest epiphytes (e.g.Dicranoweisia cirrata, Dicranum tauricum, and Scapania bolanderi), or on other substrata inthe boulder slope, especially on soil (e.g. Dicranum scoparium and Pohlia nutans), or on bothwood and soil (e.g. Rhytidiadelphus triquetrus and Jamesoniella autumnalis). In terms ofmoisture conditions, wood and soil in the boulder slope and trees in the forest could be seenas middle grounds between the extreme of forest and boulder slope. Trunks and branches oftrees where these species are usually found in the forest are likely to be among the driest offorest microhabitats. On the other hand, soil and wood are likely to be among themicrohabitats of the boulder slope which retain moisture the longest.CliffsThe results have shown that cliff habitats are markedly diversified in terms ofmicroenvironmental conditions. This was reflected by a relatively high species diversity75(Table 3.1) and a large number of species shared with nearly all of the other habitats (Table3.4, Fig. 3.1). As mentioned earlier, a key factor to species diversity is likely to be the varietyof moisture conditions found along the cliffs. In very wet sites are found species likeCratoneuron filicinum, Fissidens grandifrons, Hymenostylium insigne, and Palustriellacommutata. Moist sites harboured species such as Campylium stellatum, Didymodon rigidulusvar. gracilis, Hymenostylium recurvirostre, Blepharostoma trichophyllum, Moerckia hibernica,and PeIlia neesiana. In more mesic situations, we encountered species such as Anomolnyumfiliforme, Ditrichum crispatissimum, Encalypta spp., Mnium thomsonii, Plagiopus oederiana,Seligeria donniana, S. tristichoides, TorteIla tortuosa, Jun germannia atrovirens, Preissiaquadrata, and Reboulia hemisphaerica. Finally, in dry sites, grew species such asHeterocladium macounii, Racomitrium aciculare, Schistidium apocarpum var. apocarpum,Frullania caltfornica, and Scapania americana. It is in the two latter sites that occurred alarge proportion of the species shared with the forest and the boulder slope (examples of thesespecies were given earlier in the text). Light conditions along the cliffs are also variable.Although all cliffs have more or less the same northwestern exposure, some are open whileothers are shaded to varying degrees by the adjacent forest. Finally, although most rocksurfaces along the cliffs are calcareous, non-calcareous rock is also present. The differencebetween rock types is not only chemical. Different types of rock also have different water-holding capacity (Smith 1982). For instance, limestone holds more water than siliceous rock.Several species found on cliffs were closely associated with very particularcombinations of conditions. In many cases, although conditions between two sites seemedsuperfic ially similar, bryophyte communities differed. For instance, species such as76Anomobiyum fihforme, Seligeria donniana and S. tristichoides, and Pellia neesiana werefound only at very specific sites along the cliffs, while their microhabitat seems repeated inseveral places.ForestLike the boulder slope, the forest contained a large number of species, as well as ahigh proportion of species unique to it. Again, such results can be attributed to environmentalconditions unique to the forest. As seen in Figure 3.1, of all the species occurring onvegetation throughout the study area, the majority (94%) were forest species. It is a similarsituation for a large proportion of the species occurring on wood (62%). In other words,epiphytes and species growing on wood are best represented in the forest. From such resultswe can conclude that abundance of vegetation, and consequently abundance of wood, assubstrata is particular to the forest. Several of the species found only in the forest areassociated with trees or wood, for example: Bwcbaumia piperi, Dicranum fuscescens, Hypnumcircinale, Neckera pennata, Orthotri chum spp., Plagiothecium undulatum, Ulota spp.,Calypogeia fissa, Lepidozia reptans, Metzgeria temperata, and Porella cordaeana. It is ofinterest to note that most of these species can grow on rock surfaces elsewhere.Considerable variation exists in the composition of bryophyte communities on treesand on wood. In the case of trees, the variation observed has been related to a series ofinteracting factors which influence light conditions, relative humidity, nutrient availability andthe chemical and physical nature of the substratum. These factors include the age of the hosttree, height on the tree, direction of exposure, inclination, and chemical and physical77characteristics of the bark, such as porosity, roughness, exfoliation, chemistry, pH. (Slack1976, 1977; Smith 1982; Kenkel and Bradfield 1981, 1986; and Schofield 1985). Such a largenumber of interacting factors makes it difficult to discern clear patterns in the distribution ofepiphytic species in the results of this study. This would require precise measurements ofmicrohabitat characteristics and systematic sampling of epiphytic communities. Nevertheless,some trends are apparent. For example, desiccation-tolerant species, such as Dicranoweisiacirrata, Orthotrichum spp., and Tortula ruralis, were found in the canopy of the trees wherethere is higher illumination and lower relative humidity. On the other hand, at the base oftrees we found species associated with rich, moist habitats, such as Plagiothecium undulatum,PseudotaxiPhyllum elegans, and Rhizomnium glabrescens.As pointed out by Slack (1976), the nature of epiphytic communities is dynamic. As atree grows, surrounding environmental conditions are modified, and the nature of the barkchanges. This results in changes in the structure and composition of epiphytic communitiesthroughout the life of the tree. After the tree falls, moisture and illumination of the treechanges and the epiphytic communities which occupied the living tree are replaced by others(Schofield 1985). These new communities are in turn replaced as the wood decays and as thesurrounding vegetation responds to the local disturbance. For instance, we found species ofOrthotrichum only on recently-fallen wood. Other species, like Leucolepis acanthoneuron,Metzgeria temperata and Scapania americana, occurred on wood in an early stage of decay.Finally, some species were associated with rotten wood. Examples are BuxbaumiaPlagiothecium laetum, Pseudotaxiphyllum elegans, Tetraphis pellucida, Calypogeia fissa, andLepidozia reptans.78Although the forest harboured a large number of epiphytes, only a small number ofspecies were restricted to trees. This corroborates results presented by Smith (1982) and Slack(1977). Results presented by Smith (1982) concerning British mosses showed that themajority of epiphytes are not restricted to epiphytic conditions. In the British flora, there are33 obligate epiphytic mosses versus 44 facultative ones. Similarly, Slack (1977), working inforest stands on Ampersand Mountains in the Adirondack, found that only nine of 36epiphytic bryophyte species grew exclusively on trees. Similarly, few of the numerous speciesgrowing on wood are restricted to that substratum. In fact, several forest species occur bothon wood and on living trees. The index of similarity between forest vegetation and forestwood is relatively high (0.5). Examples of species growing both as epiphytes and on wood(apart from species found on recently-fallen wood) are: Dicranum fuscescens, Hypnumcircinale, Isothecium myosuroides, Plagiothecium undulatum, Rhizomnium glabrecens,Lophocolea cuspidata, Metzgeria conjugata, M. temperata, and Scapania bolanderi.Furthermore an important proportion of forest epiphytes are also forest floor species. Theindex of similarity between the two latter habitats was 0.23. When on trees, the speciesinvolved were often found growing at tree bases. Examples are Antitrichia califomica,Eurhynchium oreganum, E. praelon gum, Leucolepis acanthoneuron, Plagiomnium insigne, P.rostratum, Rhytidiadelphus loreus and Thamnobryum neckeroides. Several of the latter speciescan grow also on wood. Therefore, our results also show a relatively high degree of similaritybetween the forest floor flora and the flora of forest wood (0.25).79Streams and spray zonesAs described above, stream and spray zone habitats both had low species diversity. Ithas been related to their small area. This low species diversity could also be related to thehigher degree of specificity associated with habitats such as streams and spray zones. Theenvironmental conditions found in these habitats can be viewed as somewhat restrictive. Slackand Glime (1985), for instance, describe stream habitats as unstable. They note that only avery small proportion of species can establish themselves and survive under streamconditions. They pointed out the problems associated with withstanding currents and abrasion,and survival of repeated desiccation and flooding according to varying water levels. To theserestrictive conditions, we could add the accumulation of sediments, and more particularly tothe study area, the water's high concentration of calcium. With the exception of currents andabrasion, all these conditions are found also in spray zones.Nevertheless, despite the certain degree of specificity which can be attributed to thesetwo habitats, streams and spray zones had only a relatively low proportion of unique species.Species unique to stream habitats are Brotherella roellii, Campylium chrysophyllum,Platyhypnidium riparioides, Porotrichum bigelovii, and Chiloscyphus polyanthos. Speciesfound only in spray zone habitats are Dicranella varia, Eurhynchium pulchellum var. barnesii,Jun germannia leiantha, and Riccardia chamedryfolia. As described above, this low proportionof unique species partly reflects the fact that several species were shared with high-moisturehabitats found along the cliffs. However, it could be related also to an edge effect. Thehabitat categories that were defined for this study overlap to a certain degree. This is trueespecially as edges between habitats provide environmental conditions intermediate between80those characteristic of each habitat. The effect of edges on habitat categorization isparticularly important in smaller habitats, such as streams and spray zones, where theproportion of edges to the bulk of the habitat is much greater. It is even more pronounced ina long and narrow type of habitat such as streams. The precise limits of stream banks, forinstance, are not clear. A similar situation prevails in the extent of spray zones. The transitionbetween stream, or spray zone habitats, and forest habitats is not sudden. It involves gradualchanges in environmental conditions, particularly relative humidity. There is also a temporalelement involved, dictated by fluctuations in water levels. Spray zones, for instance, are widerduring periods of higher precipitation.3.4. ConclusionThe results of this study provide the basis for a general discussion of the ecology ofbryophytes at Bridal Veil Falls. Although discussion of these results involves considerablespeculation, it has raised a series of hypotheses and highlighted trends that could serve as abasis for more detailed ecological work.General differences between habitats in terms of environmental conditions were foundto play a role as a promotor of overall species diversity at the site. Such differences involvedexposure, light and moisture conditions, edaphic conditions, and type of substratum dominantin the habitat. At a smaller scale, habitat complexity was found to be a source of diversity. Awide variety of microhabitats available to bryophytes characterized the most diverse habitats.Involved here were again light, moisture and edaphic conditions, as well as factors such asmicrotopography and cyclic successional changes.81Chapter 4. Phytogeography4.1. Materials and methods4.1.1. Definition of phytogeographic elementsThe bryophyte species of Bridal Veil Falls were grouped by affinities between theirdistributions. The approach used, based upon the recognition of phytogeographic elements,has been described by Stott (1981). Phytogeographic elements were defined in terms of thedistribution of the bryophytes in the Northern Hemisphere, north of the tropics (Table 4.1). Asecond set of phytogeographic elements was used to describe their regional distributionswithin Pacific North America (Table 4.2). The Pacific North American region includes thePacific coast of Alaska, and the regions west of the Rockies in Yukon, British Columbia andthe United States.Northern hemisphere distributions of the various species were determined primarily onthe basis of distributional information provided by Nyholm (1975), Ireland (1982), and Smith(1978) for the mosses; and by Godfrey (1977) for the liverworts. Regional distributions of themosses were assessed based primarily upon distributional information provided by Vitt et. al.(1988), Lawton (1971), and Schofield (unpublished data). Taxonomic treatments byHoisington (1979), Vitt (1973), Frisvoll (1983, 1988), and Ireland (1970) provided additionalsources of distributional information. Regional distributions of liverworts were derived fromVitt et. al. (1988), Schofield (unpublished data), Godfrey (1977), and Schuster (1966, 1969,1974, 1980, 1992a, and 1992b). In addition, taxonomic treatments by Hong (1980, 1992) andHong eta (1990) were also useful.82Table 4.1. Phytogeographic elements based on distributions within the Northern Hemisphere.(modified from Godfrey 1977)I. Continuous distributionsA. Circumboreal distributions1. Circumboreal (CB)2. Circumboreal, missing in Eastern Asia (CB(-EA))3. Arctic-alpine (AA)B. Endemic distributions1. Endemic to Western North America (WNA)2. Endemic to North America (NA)II. Discontinuous, or disjunct, distributionsA. Disjuncts of european affinities1. North America/Western Eurasia (NA/WE)2. Western North America/Western Eurasia (WNA/WE)3. Western North America/Newfoundland and/or Greenland/Western Europe(WNA/ENA/WE)B. Disjuncts of eastern asian affinities1. North America/Japan (NA/J)2. Western North America/Eastern Asia (WNA/EA)C. Disjuncts of mixed affinities1. Western North America/Western Eurasia/Eastern Asia (WNA/WE/EA)2. Western North America/Eastern North America/Western Eurasia/EasternAsia (WNA/ENA/WE/EA)83Table 4.2. Phytogeographic elements based on regional (Pacific North American)distributions.I. Northern Distributions - with southern British Columbia, or northern Washingtonpopulations marking the southern limit of the range1. Widespread (nw)2. Coastal - restricted to the coast (nc)3. Discontinuous - restricted to the coast and humid interior (nd)II. Southern Distributions - with southern British Columbia populations marking the northernlimit of the range1. Widespread (sw)2. Coastal - restricted to the mediterranean (winter wet/summer dry) climate of thecoast (sc)3. Discontinuous - restricted to the mediterranean climate (winter wet/summer dry) ofthe coast and dry sites of the interior (sd)III. Widespread Distributions - with southern British Columbia populations more or less in themiddle of the range1. Widespread (w)2. Coastal - restricted to the coast (c)3. Discontinuous - restricted to the coast and humid interior (d)IV. Arctic-montane Distributions - of arctic/alpine or subarctic/subalpine affinity1. Arctic-montane (am)844.2. Results and discussion4.2.1. Affinities of the flora - broad geographic perspective.Twelve phytogeographical elements were used to describe the Northern Hemisphericrange of the bryophytes of Bridal Veil Falls (Table 4.1). The element to which each specieswas assigned is given in Appendices A and B. Table 4.3 summarizes the elements representedat Bridal Veil Falls. The following presents a general discussion of these variousdistributional elements. Predominant patterns are discussed. The detailed discussion ofparticular cases is beyond the scope of this study.Most of Bridal Veil Falls' flora (60%) is circumboreal (Table 4.3, elements 1 to 3).The circumboreal elements include species that have a more or less continuous distributionthroughout the Northern Hemisphere (e.g. Ceratodon purpureus, Climacium dendroides,Cratoneuron filicinum, Dicranum scoparium, Funaria hygrometrica, Hylocomium splendens(Figure 4.1), Neckera pennata, Plagiothecium denticulatum, Sanionia uncinata, Tetraphispellucida, Conocephalum conicum, and Lepidozia reptans). Circumboreal species dominatethe bryophyte flora of the holarctic. The large number of species showing such a distributionis attributable to a number of factors (Godfrey 1977; Schofield 1980, 1992). The ice sheetsthat spread over large portions of the northern continents during the Pleistocene shaped asimilar landscapes. After their retreat, they left vast tracts of land available for colonization.The large number of circumboreal taxa reflects also climatic uniformity throughout much ofthe Northern Hemisphere. Finally, the land masses involved are relatively continuous atpresent and have been since long in the past. As noted in Godfrey (1977), extended periods ofgeographical contact or proximity provided the opportunities for dispersal.85Table 4.3. The number of bryophytes belonging to each circumboreal phytogeographicelements.Abbreviations of phytogeographic elements is as follows: AA=arctic-alpine,CB=circumboreal, EA=eastern Asia, ENA=eastern North America, J=Japan, NA=NorthAmerica, WE=western eurasia, WNA=western North America. (see Table 4.1)PhytogeographicelementsNumber ofmossesNumber ofliverwortsTotal Percent ofthe flora(Y.")1. CB 88 31 119 562. CB(-EA) 0 4 4 23. AA 0 4 4 24. WNA 29 5 34 165. NA 3 1 4 26. NA/WE 9 3 12 67. WNA/WE 13 1 14 78. WNA/ENANVE 1 2 3 19. NA/J 1 2 3 110. WNA/EA 4 3 7 311. WNA/WE/EA 2 0 2 112. WNA/ENA/WE/EA 2 4 6 386Figure 4.1. World distribution of Hylocomium splendens. (from Schofield 1974).87In the case of arctic-alpine species, although their distribution is continuous, theypredominate at high northern latitudes, extending southward in higher elevations of mountainsor in other microenvironments that closely match the macroenvironment of arctic regions(Schofield 1969a). For instance, the extension of arctic bryoflora to western Newfoundlandcoincides with the southerly flow of cold Labrador ocean current that produces cold summers(Schofield 1992). Examples of arctic-alpine species at Bridal Veil Falls are Lophozia alpestrisand Tritomaria quinquedentata.Twenty-two percent of the flora is of species that have discontinuous distributionsthroughout the Northern Hemisphere (Table 4.3, elements 6 to 12). These species show widegaps, or disjunctions, in their range. Large continental disjunctions are generally thought of asremnants of more continuous distributions of the past. Most disjunct species are restricted tovery specific climatic regions that have probably been interconnected in the very distant pastand are markedly disjunctive at present. Such disjunctions thus originate over extensivegeological time periods - the results of various historical factors such as changes in thecontinental positions or variation in climate through time (Belland 1987; Schofield 1985,1988; Schofield and Crum 1972). In most cases, it is unlikely that long distance dispersal wasinvolved, despite the fact that most bryophytes have diaspores highly suitable for longdistance wind-dispersal. The disjunctive distribution patterns observed in bryophytes coincidemostly to the same patterns observed in vascular plants. It is thus likely that disjunctions ofvascular plants and bryophytes followed the same historical development (Schofield 1980).Furthermore, a high proportion of disjunctive species are dioicous, and do not produce anyspecialized vegetative propagules for dissemination (Schofield 1980, 1984; Schofield and88Crum 1972). Disjunctive species are also often associated with highly humid oceanic climates,often in mountainous regions where the effects of strong winds and the opportunities forpropagules to become air-borne are considerably reduced (Schofield 1980; Schofield andCrum 1972; Schuster 1983). This is especially true of species that have long-establishedassociation with forest vegetation (Schofield 1984). Furthermore, disjunctive species are foundmostly in or near regions that escaped the harshest extremes of the glaciation (Schofield1988).At Bridal Veil Falls, species of the discontinuous elements are usually disjunct to oneor more of the following regions: western North America, eastern North America, easternAsia, and western Eurasia. A large proportion are disjuncts between western North Americaand western Eurasia. These western North American-western Eurasian disjuncts represent twomain types of disjunction as described in Schofield (1985, 1988). The first type involvesdisjunctive species which occur in the oceanic (or highly humid) climates of western NorthAmerica and western Europe (e.g. Dicranoweisia cirrata, Dicranum tauricum, Hymenostyliuminsigne (Figure 4.2), Plagiothecium undulatum, and Porella cordaeana). In western NorthAmerica, oceanic climates are found eastward of the mountain chains, predominantly thecoastal ones, which act in generating moist climates under the influence of dominant westerlywinds. The greatest diversity of oceanic species is in northern coastal regions of Pacific NorthAmerica. The second type of western Eurasian-western North American disjunction involvesspecies found in the mediterranean (or winter wet, summer dry) climates of the southernportion of the coast of western North America and the coastal regions of the Mediterranean inEurope and northern Africa (e.g. Antitrichia cahfornica (Figure 4.3), Metaneckera menziesii,89Figure 4.2. World distribution of Hymenostylium insigne. (modified from Schofield 1989).90Figure 4.3. World distribution of Antitrichia califomica. (from Schofield 1988).91Orthotrichum lyellii, Scleropodium cespitans, and Tortula princeps). In western NorthAmerica, the range of mediterranean species can extend from southern California to southernBritish Columbia, and in some cases northward as far as southwestern Alaska in suitable,drier microclimates. Some of the species can occur also inland in edaphically favourable areas(Schofield 1980).Oceanic climates, as those described above, tend to be particularly rich in disjunctivetaxa (Schofield and Crum 1972). Such climates occur not only in Pacific North America andwestern Eurasia but also in eastern Asia. Disjuncts are common also between oceanic regionsof western North America and eastern Asia (Hong 1987; Schofield 1965; and Schofield andCrum 1972). Among the western North American-eastern Asian disjuncts found at Bridal VeilFalls are Claopodium crispifolium (Figure 4.4a), Hypnum subimponens (Figure 4.5a),Oligotrichum aligerum, and Pogonatum contortum. In a few cases, oceanic disjuncts occuralso in eastern North America. Examples are Antitrichia curtipendula, Bryum miniatum,Pseudotaxiphyllum elegans (Figure 4.6a), Rhytidiadelphus loreus, and Tetraphis geniculata.As seen above, most disjunctive species are environmentally restricted (Schofield1969a). However, in a few cases, disjunct species do not seem associated with a well-definedset of environmental conditions. Examples are Epipterygium tozeri, Grimmia torquata, andTortula subulata. Epipterygium tozeri, for instance, occurs not only in meditteffanean climatesin western North America and western Eurasia, but extends to different climates in southeastAsia and Britain. The disjunct distribution of such species could represent fragments of acircumboreal Tertiary flora (Schofield 1965, 1980).92Figure 4.4. Claopodium crispifolium, a) world distribution (modified from Schofield 1965),b) Pacific North American distribution (from Schofield, unpublished data).B)93Figure 4.5. Hypnum subimponens, a) world distribution (from Schofield 1965), b) PacificNorth American distribution (from Schofield, unpublished data).A)B)94Figure 4.6. Pseudotaxiphyllum elegans, a) world distribution (data from BeHand 1987, andStOrmer 1969), b) Pacific North American distribution (from Schofield, unpublished data).B)95Finally, eighteen percent of the bryophytes at Bridal Veil Falls are restricted to NorthAmerica (Table 4.3, elements 4 and 5). The majority are western North American endemics.These are confined mainly to areas in, or west of, the Rocky Mountains (Schofield 1969a).Like the western North American-western Eurasian disjuncts, western North Americanendemics found at Bridal Veil Falls can be of two types: 1) oceanic, associated with highlyhumid climates found primarily west of the mountain ranges of the Cordillera (e.g.Buxbaumia piperi, Eurhynchium ore ganum, Leucolepis acanthoneuron, Plagiomniumvenustum, Racomitrium lawtonae (Figure 4.7), Thamnobryum neckeroides, PoreIlanavicularis, and Scapania americana), or 2), mediterranean, associated with the moresouthern, winter wet, summer dry climates (e.g. Amphidium californicum, Anacolia menziesii,Dendroalsia abietina, Pseudobraunia californica (Figure 4.8), Racomitrium pacificum, andFrullania californica). In both cases some species are restricted to the coast, while othersoccur also inland in suitable microclimates. According to Schofield (1980, 1984), the bulk ofthis western North American endemic flora , like the disjunctive flora of the region, wouldhave survived the glaciations south of the glacial boundary in the western United States.A small proportion of the species that are confined to North America are found alsooutside of Pacific North America. In most cases, these species are disjuncts between westernand eastern North America. As discussed in BeBand (1987) and Belland and Schofield (1988),the absence of many North American species in the central portion of the continent coincideswith the presence of the prairie biome. There are several examples of species that showdisjunctions between western and eastern North America, and are not found in the GreatPlains. Examples of such species at Bridal Veil Falls are: Grimmia torquata, Heterocladium96504030Figure 4.7. Pacific North American distribution of the endemic Racomitrium lawtonae (fromFrisvoll 1988, and Schofield, unpublished data). (Also known from the Aleutians)97Figure 4.8. Pacific North American distribution of the endemic Pseudobraunia californica(from Schofield, unpublished data).98macounii, Pseudoleskea stenophylla, Pseudotaxiphyllum elegans (Figure 4.6a),Rhytidiadelphus loreus, Tetraphis geniculata, and Frullania tamarisci subsp. nisquallensis.In summary, at Bridal Veil Falls, the distribution of species between broadphytogeographic elements is typical of the bryophyte flora of northwestern North America asdescribed by Schofield (1968b, 1968c, 1969a, 1984 and 1988). The bulk of the flora iscircumboreal. It also is rich in endemics. Finally, it is characterized by the presence of widedisjuncts, usually to western Eurasia and to eastern Asia, and predominantly associated withoceanic environments.4.2.2. Affinities of the flora - regional geographic perspective.Ten phytogeographical elements were defined (Table 4.2) to describe the Pacific NorthAmerican distributions of the species collected at Bridal Veil Falls. The phytogeographicalelement to which each species was assigned is indicated in Appendices A and B. Table 4.4provides a summary. Two percent of the species remained unclassified either because theirdistribution was poorly understood or because it did not accommodate any of thedistributional elements defined and it seemed superfluous to create categories to accommodatethem individually.The majority of the species found at Bridal Veil Falls (55%) are widespreadthroughout Pacific North America. Most to these species have circumboreal distribution.Also representing an important percentage of the flora are widespread discontinuousand widespread coastal species with 20% and 10% of the flora respectively. These are mostlyendemic or disjunctive species associated with the humid oceanic climates of western North99Table 4.4. The number of bryophytes belonging to each regional phytogeographic elements.Phytogeographic elements NumberofmossesNumber ofliverwortsTotal Percent ofthe flora(%)Widespread 89 27 116 55Widespread coastal 8 14 22 10Widespread discontinuous 32 10 42 20Arctic-montane 2 5 7 3Southern widespread 0 1 1 0.5Southern coastal 4 2 6 3Southern discontinuous 7 0 7 3Northern widespread 4 0 4 2Northern coastal 1 0 2 0.5Northern discontinuous 2 0 1 1Unclassified 3 1 4 2100America. Coastal species are restricted to the humid coastal regions (e.g. Claopodiumcrispifolium (Figure 4.4b)), while discontinuous species occur as well into the interior, mainlyin the Columbia Mountain ranges (e.g. Hypnum subimponens (Figure 4.5b) andPseudotaxiphyllum elegans (Figure 4.6b)). The Columbia region shows many similarities tothe coast in terms of climate and vegetation (Schofield 1976, 1988). Easterly moving airmasses produces cool wet winters and warm dry summers, and as in the coastal forest, thetree species that dominate in mature stands are Tsuga heterophylla and Thuja plicata(Ketcheson et al. 1991).The remaining 13% of the flora consists of species of northern, southern, or arctic-montane distributions. The Bridal Veil Falls populations of these species are at the limit oftheir range, or in the case of arctic-montane species, represent unusual extension of the range.Northern coastal or northern discontinuous species are, as above, endemic or disjunct speciesassociated with the humid climates of the coast and the interior. In this case, however, theyhave a generally more northern distribution. Examples are Hymenostylium insigne (Figure4.2), Racomitrium lawtonae (Figure 4.7), and Tetraphis geniculata. Unlike the morewidespread coastal or discontinuous species which are probably immigrants from populationsthat survived south of the glacial boundary, these more northern species are thought to bedescendants of relictual populations in unglaciated areas along the coast (Schofield 1976).Southern coastal or southern discontinuous species are also mostly endemics ordisjuncts. These, however, are associated with mediterranean climates. Examples areAmphidium californicum, Anacolia menziesii, Pseudobraunia californica (Figure 4.8), andFrullania califomica.101Northern and southern widespread species are species that are more widely distributedin the holarctic. In the case of northern ones, their range is restricted to northern temperateand/or boreal regions (e.g. Polytri chum formosum, Seligeria campylopoda, and S. donniana).On the other hand, southern widespread species are restricted to southern temperate regions(e.g. Reboulia hemisphaerica).4.2.3. Relationship between habitat and geographic affinity.This section examines the relationship between the Pacific North American distributionof species and their occurrence in the various types of habitat available at Bridal Veil Falls.Table 4.5 gives details as to how the different regional phytogeographic elements arerepresented in each habitat. The distribution of widespread species between habitatscorresponds to the way the whole of the Bridal Veil Falls' flora is generally distributedbetween habitats (see Table 3.1). Diversity of widespread species is higher in the habitatswith higher species diversity and lower in the ones with lower species diversity. On the otherhand, although coastal and discontinuous species are generally more numerous in the morediverse habitats, they are the most numerous in the forest. Such results could be related to themore even character of forest habitats. Climatic conditions in forest are not subject to asmuch fluctuation as in more open habitats such as the boulder slope. The high humidity ofcoastal regions which is favourable to growth of bryophytes is maintained more uniformlyunder forest conditions. The different types of microhabitats available to bryophytes in forestenvironments are also less variable than in other habitats like cliffs or boulder slope. Thelatter can show considerable variation in terms of edaphic conditions, exposure, humidity,102Table 4.5. Number of bryophytes species of each regional phytogeographic element occurringin each habitat type.Abbreviation of phytogeographic elements is as follows: am=arctic-montane, c=coastal,d=discontinuous, n=northern, s=southern, and w=widespread. (see Table 4.2)Habitat types Number of species of each affinityW C D NW NC ND SW SC SD AM UAnthropogenic 15 1 4 0 0 0 0 0 0 0 0Boulder slope 44 5 20 0 0 1 1 0 3 3 0Cliff 49 6 15 2 1 0 1 1 1 2 0Forest 41 10 31 1 0 0 0 2 2 0 1Stream 23 2 7 0 0 0 0 0 2 0 1Spray zones 23 5 4 0 1 0 0 0 0 0 0Note: This table includes only data on the species collected at Bridal Veil Falls during the course of thisstudy.103etc. Humid forest habitats hence form a more uniform unit through space and time allowingthe wider dispersal of the bryophytes associated with them.Godfrey (1977) examined regional distribution patterns of hepatics in southwesternBritish Columbia. She observed that regional distribution patterns appear to reflect patterns ofavailability of particular microclimatic conditions. Therefore, the wide variety ofmicroenvironments available at a particular site would not only promote species diversity (seeChapter 3), but it could also promote the diversity of phytogeographic elements represented atthe site. As stated in Godfrey (1977), suitable microclimate can affect regional distribution inextending the local opportunities for occurrence of species of different affinities at a site.Bryophytes near the limit of their range are particularly susceptible to the availabilityof particular microclimatic conditions. Reporting BOcher (1954), Hedderson (1987) remarkedon the importance of microhabitats for bryophytes, especially near the limits of speciesdistributions, where plants depend more strongly on definite and limited combinations ofmicrohabitat factors. Different authors (BeHand and Brassard 1988; Hedderson 1987; Vaaramaand Laine 1974) have observed how species at the limit of their range in their respectivestudy areas were restricted in terms of habitat to a narrow range of favourable microhabitatconditions. At Bridal Veil Falls, bryophytes near the limit of their range include species ofnorthern, southern and arctic-montane distribution. As expected, these species are oftennarrowly restricted in terms of the habitats and substrata on which they were found; several ofthem are among the species that were found confined to one habitat or substratum.In the previous chapter, boulder slope, cliff and forest habitats were identified as themost diverse in terms of microenvironments available to bryophytes. It is interesting to note104that it is in these three habitats that species of northern, southern and arctic-montanedistribution are mainly found (Table 4.5). This could be simply an artefact of the largenumber of species found in these habitats. Nevertheless, the wide variety ofmicroenvironments in these habitats is likely to be also an important factor considering theimportance of particular microhabitats for the establishment and survival of bryophytes at thelimit of their range. If a wide variety of microhabitats are available at a site, it is more likelythat this site will offer suitable conditions.Godfrey (1977) classified the liverworts she collected in southwestern British columbiaaccording to patterns of local distribution for the region. Among the species of liverwortsfound at Bridal Veil Falls are species that she classified as strictly oceanic: Chandonanthusfiliformis, Moerckia hibernica, and Odontochisma denudatum. She described these species asrestricted to coasts along the Pacific Ocean, occurring along the west coast of VancouverIsland and northward along mainland inlets. These species were said to be rare or absent inthe Lower Mainland. Their local distribution is shown in figure 4.9. Although Godfrey (1977)was not aware of the presence of these species at Bridal Veil Falls, this would probably nothave changed their strictly oceanic status. Bridal Veil Falls' populations are isolatedpopulations of these species and their presence inland remains exceptional. A few species ofmosses could also be classified as strictly oceanic. Among the mosses found at Bridal VeilFalls, the only truly strictly oceanic species is Hymenostylium insigne which is knownelsewhere in western North America only from the Queen Charlotte Islands (Figure 4.2). Thepresence of these strictly oceanic species inland at Bridal Veil Falls is unusual. It suggests theavailability of microhabitats where humidity is exceptionally high for the region.105Figure 4.9. Regional distribution of a) Chandonanthus fihformis, b) Moerckia hibernica, andc) Odontochisma denudatum.A) Chandonanthus filiformisB) Moerckia hibemicaC) Odontochisma denudatum106Odontochisma denudatum, for example, was found on a rotten log high in the boulderslope. The site was shaded by high cliffs, thus moisture persisted there most of the year.Similarly, Hymenostylium insigne and Moerckia hibernica were both associated with dampcliff faces, usually in proximity of waterfalls. Chandonathus filiformis, although on a dry rocksurface, was found in a shaded and sheltered site. In general, the relatively cooler and moremesic climate characterizing Bridal Veil Falls (see Chapter 1) is likely to be an importantfactor accounting for the presence of the above species. Availability of calcareous substrata isalso a factor for calcicoles such as Moerckia hybernica and Hymenostylium insigne.The presence of arctic-montane species at Bridal Veil Falls is also unusual (e.g.Barbula convoluta var. gallinula, Plagiobryum zierii, Anastrophyllum assimile, Lophoziaalpestris, and Tritomaria quinquedentata). Such species were described by Godfrey (1977) assubalpine-descending. In other words, these species are chiefly high elevation, butoccasionally descend to low elevations in microclimatically suitable sites. Godfrey mentionedwaterfalls and cold, humid boulder slopes or rock outcrop-slopes as examples of such sites.As expected, arctic-montane species at Bridal Veil Falls were associated with north-facingcliffs and the boulder slope (Table 4.5). The availability of cold, humid microsites along thecliffs has already been mentioned. It is in such a site that the arctic-montane species Barbulaconvoluta var. gallinula and Plagiobryum zierii were found. Both species were found inadjacent sites on a moist, shaded cliff. In the case of the boulder slope, the emphasisthroughout the text has been that it was an open, dry and warm site. It could, however, alsobe described as a cold, humid habitat. In fact, there is a seasonal pattern in the microclimaticconditions characterizing the boulder slope. During the fall and winter, precipitation is greater107(Figure 1.5), and duration of direct sunlight is considerably reduced, even more so consideringthe high elevation of the adjacent cliffs; the boulder slope thus becomes a cold, humid habitat.For example, the presence of Polypodium glycyrrhiza in the boulder slope is not noticeable inthe summer. By late fall and throughout winter, this fern flourishes in masses over theboulders. Such cold, humid conditions as described above are likely to lag behind, as summerapproaches, in microsites on the north sides of boulder and closer to the cliff bases.Furthermore, if snow accumulates during the winter, it is likely to stay longer in thesemicrosites keeping them cooler. Arctic-montane bryophytes are usually associated with thesecolder, moister microsites in the boulder slope (e.g. Anastrophyllum assimile).Also found at Bridal Veil Falls is a group of species that Godfrey (1977) classified asspecies of dry southern habitats. This group includes most species described earlier assouthern (e.g. Frullania bolanderi, F. cahfornica, and Reboulia hemisphaerica). Godfrey(1977) observed that these species generally occupy dry, relatively sunny areas. Althoughspecific requirements vary between species, these dry, southern species usually do, in fact,occupy drier and warmer habitats at Bridal Veil Falls. Those that occur as epiphytes werefound on trunks, branches, and in the canopy of trees rather than at tree bases whereconditions are presumably moister (e.g. Antitrichia cahfornica, Dendroalsia abietina, Tortulasubulata, and Frullania bolanderi). Species occurring on cliffs were associated with dry rocksurfaces (e.g. Frullania cahfornica). Finally, dry southern species occurring in the boulderslope (e.g. Amphidium californicum, Anacolia menziesii, Pseudobraunia cahfornica, andReboulia hemisphaerica) were found mostly on open, exposed sites or on dry boulder faces.1084.2.4. An interpretation of the development of the flora.As seen above, the flora of Bridal Veil Falls contains a number of species that arephytogeographically significant. These would include species that are closely associated withvery specific microenvironmental conditions found at the site, rare species, and species thatshow important gaps in their regional distribution. High species diversity at the site andpresence of numerous phytogeographically significant species raises questions as to the originand development of the bryoflora.Considering the late-Pleistocene and Holocene history of the area, the flora of BridalVeil Falls is relatively recent. The vegetation that colonized and flourished at the sitefollowing the retreat of the Sumas glacier at the end of the last glaciation was probablycompletely destroyed as a result of the Cheam Slide. Subsequent plant communities thatestablished themselves at the site were in turn disturbed by more recent events, such aslogging and rockfall, as well as smaller scale events, such as the installation of a pipeline andof park facilities. As seen in chapter three, these recent events have been mostly promoters ofbryophyte diversity as they enhanced the variety of habitats available for bryophytes.The bulk of the bryoflora of Bridal Veil Falls has its source in adjacent areas of theFraser Lowland. Bryophytes from these adjacent areas were undoubtedly the main source ofpropagules for colonization after disturbances. As seen earlier, the flora of the region ispredominantly of circumboreal affinity, with a strong humid coastal component. It probablyderives primarily from bryophyte populations that survived the Wisconsin glaciation south ofthe glacial boundary. In a few cases, floristic elements could possibly derive from bryophytepopulations that survived in coastal refugia along the coast of southern British Columbia, or109in mountain refugia in adjacent regions (Godfrey 1977). Some species could have invaded thearea preceding the establishment of the modern vascular plant vegetation, while othersprobably moved in with the latter and depend on its presence for survival. Acermacrophyllum, for example, a tree that harbours abundant and diverse bryophyte communities,is likely to be a late-comer in the Fraser Lowland and at Bridal Veil Falls. It is a species ofmore southern distribution and it characterizes mostly second growth stands.The expansion of the range of mediterranean elements to the Fraser Lowland has beenassociated with the Hypsithermal period (Schofield 1988). It is possible that the southernspecies took advantage of this warmer and drier period to extend the limit of their rangenorthward, and, at the outset of that period, survived in these northern areas where favourablemicroenvironments were available. Nevertheless, as argued by Godfrey (1977), a xerothermicperiod is not necessary to explain the occurrence of species of dry southern climates insouthwestern British Columbia. Their presence might simply be the result of their movementnorthward, as any other species displaced southward during the glaciation, into any site whichoffers favourable microenvironmental conditions.While the bulk of the modern bryoflora at Bridal Veil Falls is shared with adjacentareas, some species, however, are rare or unknown from elsewhere in the Lower Mainland. Ininterpreting the phytogeography of such species, the extent of our knowledge of bryophytedistribution becomes an important limitation. The question arises as to whether a species isreally absent from an area, or whether the area simply has not been thoroughly explored yet.Apparent disjunction can be the result of scarcity of records. This is likely to be the case ofBarbula convoluta var. gallinula, for example. There is a gap in the distribution of this110species between northern areas (Alaska, Yukon, and the Northwest Territories) and southernBritish Columbia. It is unlikely that this wide regional disjunction is the result of longdistance dispersal. Firstly, this moss reproduces essentially by means of large, bulky gemmae.Secondly, there are important geographical bathers between the two disjunct parts of itsrange.The case of oceanic species is also puzzling. These species are also often rare orunknown from the Lower Mainland. Some of these species could have dispersed inland toBridal Veil Falls from nearby western coastal regions, such as Vancouver Island (e.g.Odontochisma denudatum and Moerckia hibernica). Nevertheless, several of them are notknown to produce sporophytes or gemmae locally (Anastrophyllum assimile andChandonanthus fihformis). Other species, not only do not readily produce propagules, but areunknown from nearby coastal regions. Hymenostylium insigne is an example; its only knownlocalities, apart from Bridal Veil Falls, are on the Queen Charlotte Islands. 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Canadian Journal of Botany 60: 1596-1600.120Appendix AAnnotated list of mosses at Bridal Veil Falls.Species and authority is given first, followed by UBC accession number of the voucherspecimen, distribution in the Northern Hemisphere, and distribution in the Pacific NorthAmerica. Finally, species added to the flora by virtue of this study are marked new.Distribution in the Northern Hemisphere: AA=arctic-alpine, CB=circumboreal, EA=easternAsia, ENA=eastern North America, J=Japan, NA=North America, WE=western eurasia, andWNA=western North America. (see Table 4.1)Distribution in the Pacific North America: am=arctic-montane, c=coastal, d=discontinuous,n=northern, s=southern, and w=widespread. (see Table 4.2)Amblystegium serpens (Hedw.) Schimp. in B.S.G.; b136197; CB; wAmphidium californicum (Hampe ex C.Muell.) Broth.; b136198; WNA; scAmphidium lapponicum (Hedw.) Schimp.; b136199; CB; wAnacolia menziesii (Turn.) Par.; b136200; WNA; sdAnomobryum filiforme (Dicks.) Solms in Rabenh.; b136201; CB; wAntitrichia caltfomica Sull. in Lesq.; b136202; WNA/WE; sdAntitrichia curtipendula (Hedw.) Brid.; b136203; WNA/ENA/WE/EA; dAtrichum selwynii Aust.; b136329; NA; wAtrichum undulatum (Hedw.) P.Beauv.; b136204; CB; wAulocomnium androgynum (Hedw.) Schwaegr.; b136205; CB; wBarbula amplexifolia (Mitt.) Jaeg.; b136375; WNA/EA; unclassified, widely disjunct betweennorthern Canada and Alaska, southern British Columbia, and Arizona (Zander 1979)Barbula convoluta var. gallinula Zand.; b136206; WNA; am; new to British ColumbiaBartramia pomtformis Hedw.; b136207; CB; wBlindia acuta (Hedw.) Bruch & Schimp. in B.S.G.; b136208; CB; wBrachythecium asperrimum (Mitt.) Sull.; b136209; WNA; dBrachythecium frigidum (C.Muell.) Besch.; b136210; WNA; wBrachythecium plumosum (Hedw.) Schimp. in B.S.G.; b136211; CB; dBrachythecium rivulare Schimp. in B.S.G.; b136212; CB; wBrachythecium rutabulum (Hedw.) Schimp in B.S.G.; b136213; CB; w; new to Bridal VeilFallsBrachythecium velutinum (Hedw.) Schimp. in B.S.G.; b23910 ; CB; wBrotherella roellii (Ren.& Card. in Roell) Fleisch.; b136214; WNAI unclassified, local inhumid, lowland, coastal forests of southwestern British Columbia (Schofield 1976)Bryoerythrophyllum recurvirostre (Hedw.) Chen; b136215; CB; wBryum capillare Hedw.; b136216; CB; wBryum miniatum Lesq.; b136217; WNA/ENA/WE; dBryum pseudotriquetrum (Hedw.) Gaertn. et al.; b136218; CB; wBuxbaumia piperi Best; b136219; WNA; d; new to Bridal Veil Falls121Calliergonella cuspidata (Hedw.) Loeske; b136220; CB; wCampylium chrysophyllum (Brid.) J.Lange; b136221; CB; wCampylium polygamum (Schimp in B.S.G.) C.Jens.; b18181; CB; wCampylium stellatum (Hedw.) Ciens.; b136222; CB; wCeratodon purpureus (Hedw.) Brid.; b136223; CB; wClaopodium bolanderi Best; b94616; WNA; dClaopodium crisptfolium (Hook.) Ren.& Card.; b136224; WNA/EA; cClimacium dendroides (Hedw.) Web.& Mohr; b136225; CB; wCratoneuron filicinum (Hedw.) Spruce; b136226; CB; wDendroalsia abietina (Hook.) Brid.; b136227; WNA; sdDichodontium pellucidum (Hedw.) Schimp.; b136228; CB; wDicranella schreberiana (Hedw.) Hilf. ex Crum & Anderson; b136229; CB; wDicranella varia (Hedw.) Schimp.; b136230; CB; wDicranoweisia cirrata (Hedw.) Lindb. ex Milde; b136231; WNA/WE; dDicranum fuscescens Turn.; b136232; CB; wDicranum scoparium Hedw.; b136233; CB; wDicranum tauricum Sapeh.; b136234; WNA/VVE; wDidymodon fallax var. reflexus (Brid.) Zand.; b136235; CB; wDidymodon rigidulus var. gracilis (Schleich. ex Hook.& Grey.) Zand.; b136236; NA/WE; wDidymodon vinealis var. flaccidus (Bruch & Schimp in Schimp ) Zand.; b136237; WNA/VVE;dDistichium capillaceum (Hedw.) Bruch & Schimp. in B.S.G.; b41198; CB; wDitrichum crispatissimum (C.Muell.) Par.; b136238; CB; wEncalypta ciliata Hedw.; b136239; CB; wEncalypta procera Bruch; b136240; CB; wEncalypta rhaptocarpa Schwaegr.; b136241; CB; w; new to Bridal Veil FallsEpipterygium tozeri (Grey.) Lindb.; b136242; WNA/WE/EA; Sc; new to Bridal Veil FallsEurhynchium oreganum (Sull.) Jaeg.; b136243; WNA; dEurhynchium praelongum (Hedw.) Schimp. in B.S.G.; b136244; CB; dEurhynchium pulchellum var. barnesii (Ren.& Card.) Grout; b136245; WNA; cEurhynchium pulchellum var. pulchellum (Hedw.) Jenn.; b136246; CB; wFissidens bryoides Hedw.; b136247; CB; wFissidens grandifrons Brid.; b136248; CB; wFunaria hygrometrica Hedw.; b136249; CB; w; new to Bridal Veil FallsGrimmia torquata Hornsch. in Grey.; b136250; NA/WE; wGymnostomum aeruginosum Sm.; b136251; CB; wHedwigia ciliata (Hedw.) P.Beauv.; b20213; CB; wHeterocladium macounii Best; b136252; NA; dHomalia trichomanoides (Hedw.) Schimp. in B.S.G.; b136253; CB; wHomalothecium fulgescens (Mitt. ex C.Muell.) Lawt.; b136254; WNA; dHomalothecium nuttalii (Wils.) Jaeg.; b136255; WNA; cHygrohypnum luridum (Hedw.) Jenn.; b136256; CB; wHylocomium splendens (Hedw.) Schimp. in B.S.G.; b136257; CB; wHymenostylium insigne (Dix.) Podp.; b136258; WNA/WE; nc122Hymenostylium recurvirostre (Hedw.) Dix.; b136259; CB; wHypnum circinale Hook.; b136260; WNA; dHypnum lindbergii Mitt.; b136261; CB; w; new to Bridal Veil FallsHypnum subimponens Lesq.; b136262; WNA/EA; dIsopterygiopsis pukhella (Hedw.) Iwats.; b114080; CB; wIsothecium myosuroides Brid.; b136263; NA/WE; dLeucolepis acanthoneuron (Schwaegr.) Lindb.; b136264; WNA; dMetaneckera menziesii (Hook. in Drumm.) Steere; b136265; WNA/WE; wMnium ambiguum H.Muell.; b64819; CB; wMnium marginatum (With.) Brid. ex P.Beauv.; b136266; CB; wMnium spinulosum Bruch & Schimp. in B.S.G.; b136267; CB; wMnium thomsonii Schimp.; b136268; CB; wNeckera douglasii Hook.; b136269; WNA; dNeckera pennata Hedw.; b136270; CB; wOligotrichum aligerum Mitt.; b136271; WNA/EA; d; new to Bridal Veil FallsOrthotrichum consimile Mitt.; b136272; WNA/WE; dOrthotrichum lyellii Hook. & Tayl.; b136273; WNA/WE; dOrthotrichum pulchellum Brunt. in Winch. & Gateh.; b136274; WNA/WE; dOrthotrichum speciosum Nees in Sturm; b136275; NA/WE; wOrthotrichum striatum Hedw.; b136276; WNA/WE; dPalustriella commutata (Brid.) Ochyra; b136277; CB; wPlagiobryum zierii (Hedw.) Lindb.; b136278; CB; amPlagiomnium insigne (Mitt) T.Kop.; b136279; WNA; dPlagiomnium rostratum (Schrad.) T.Kop.; b136280; CB; wPlagiomnium venustum (Mitt.) T.Kop.; b136281; WNA; dPlagiopus oederiana (Sw.) Crum & Anderson; b136282; CB; wPlagiothecium cavtfolium (Brid.) Iwats.; b136283; CB; wPlagiothecium denticulatum (Hedw.) Schimp. in B.S.G.; b136284; CB; wPlagiothecium laetum Schimp in B.S.G.; b136285; CB; wPlagiothecium undulatum (Hedw.) Schimp. in B.S.G.; b136286; WNA/WE; dPlatydictya jungermannioides (Bud.) Crum; b136287; CB; wPlatyhypnidium riparioides (Hedw.) Dix.; b136288; CB; sdPleurozium schreberi (Brid.) Mitt.; b136289; CB; wPogonatum urnigerum (Hedw.) P.Beauv.; b136290; CB; w; new to Bridal Veil FallsPohlia cruda (Hedw.) Lindb.; b136291; CB; wPohlia nutans (Hedw.) Lindb.; b136292; CB; wPohlia wahlenbergii (Web. & Mohr) Andrews; b136293; CB; w; new to Bridal Veil FallsPolytrichastrum alpinum (Hedw.) G.L.Sm.; b136294; CB; wPolytrichum formosum Hedw.; b106819; CB; nwPolytrichum juniperinum Hedw.; b136295; CB; wPolytrichum piliferum Hedw.; b136296; CB; wPorotri chum bigelovii (Sull.) Kindb.; b136297; WNA; dPorotri chum vancouveriense (Kindb. in Mac.) Crum; b136298; WNA; c123Pottia truncata (Hedw.) Fuernr. ex B.S.G.; b84386 ; CB; unclassified, confined to essentiallycoastal agricultural areas of the Fraser River Delta (Schofield 1968b, 1976)Pseudobraunia caltfornica (Lesq.) Broth.; b136330; WNA; sdPseudoleskea stenophylla Ren. & Card. in Roe11; b94158; NA; dPseudotaxiphyllum elegans (Brid.) Iwats.; b136299; NA/WE; dPterigynandrum filtforme Hedw.; b136300; CB; wRacomitrium aciculare (Hedw.) Brid.; b136301; CB; wRacomitrium canescens (Hedw.) Brid.; b136302; CB; wRacomitrium elongatum Ehrh. ex Frisv.; b136303; NA/WE; wRacomitrium fasciculare (Hedw.) Brid.; b64806; CB; wRacomitrium heterostichum (Hedw.) Brid. sensu lato; b136304; CB; wRacomitrium lanuginosum (Hedw.) Brid.; b136305; CB; wRacomitrium lawtonae Irel.; b136306; WNA; nd; new to Bridal Veil FallsRacomitrium pacificum Irel. & Spence; b136333; WNA; scRacomitrium varium (Mitt.) Jaeg.; b136307; WNA; cRhizomnium glabrescens (Kindb.) T.Kop.; b136308; WNA; dRhytidiadelphus loreus (Hedw.) Warnst.; b136309; NA/WE; dRhytidiadelphus squarrosus (Hedw.) Warnst; b136310; CB; wRhytidiadelphus triquetrus (Hedw.) Warnst; b136311; CB; wSanionia uncinata (Hedw.) Loeske; b136312; CB; wSchistidium apocarpum var. apocarpum (Hedw.) Bruch & Schimp. in B.S.G.; b136313; CB;wSchistidium apocarpum var. strictum (Turn.) Moore; b136314; CB; wScleropodium cespitans (C.Muell.) L.Koch; b136315; WNA/WE; dScleropodium touretii var. colpophyllum (Sull.) Lawt. ex Crum; b136316; WNA; sc; new toBridal Veil FallsSeligeria campylopoda Kindb. in Mac. & Kindb.; b136317; NA/WE; nwSeligeria donniana (Sm.) C.Muell.; b136318; CB; nwSeligeria tristichoides Kindb.; b136319; NA/WE; nwSphagnum capilltfolium (Ehrh.) Hedw.; b23923; CB; wSphagnum girgensohnii Russ.; b136320; CB; wTetraphis geniculata Girg. ex Milde; b136332; NA/J; ndTetraphis pellucida Hedw.; b136321; CB; wThamnobryum neckeroides (Hook.) Lawt.; b136322; WNA; dTimmia austriaca Hedw.; b136323; CB; wTortella tortuosa (Hedw.) Limpr.; b136324; CB; wTortula princeps De Not.; b112430; WNA/WE; sdTortula ruralis (Hedw.) Gaertn. et al.; b136325; CB; wTortula subulata Hedw.; b136331; WNA/WE/EA; sdTrichodon cylindricus (Hedw.) Schimp.; b44319; WNA/ENA/WE/EA; wUlota megalospora Vent. in Roell; b136326; WNA; cUlota obtusiuscula C.Muell. & Kindb. in Mac. & Kindb.; b136327; WNA; cZygodon viridissimus (Dicks.) Brid.; b136328; CB; c124Appendix BAnnotated list of liverworts at Bridal Veil Falls.Species and authority is given first, followed by UBC accession number of the voucherspecimen, distribution in the Northern Hemisphere, and distribution in the Pacific NorthAmerica. Finally, species added to the flora by virtue of this study are marked new.Distribution in the Northern Hemisphere: AA=arctic-alpine, CB=circumboreal, EA=easternAsia, ENA=eastern North America, J=Japan, NA=North America, WE=western eurasia, andWNA=western North America. (see Table 4.1)Distribution in the Pacific North America: am=arctic-montane, c=coastal, d=discontinuous,n=northern, s=southern, and w=widespread. (see Table 4.2)Anastrophyllum assimile (Mitt.) Steph.; b136334; WNA/ENA/WE/EA; amAnastrophyllum minutum (Schreb.) Schust.; b136335; CB; wApometzgeria pubescens (Schrank) Kuwah.; b136336; CB; wBarbilophozia barbata (Schmid. ex Schreb.) Loeske; b78630; CB; wBazzania denudata (Torrey ex Gott. et al.) Trey.; b136337; NA/J; dBazzania tricrenata (Wahlenb.) Lindb.; b68270; CB; cBlepharostoma trichophyllum (L.) Dum.; b136338; CB; wCalypogeia fissa (L.) Raddi; b136339; NA/WE; unclassifiedCalypogeia trichomanis (L.) Corda; b76692; CB; dCephalozia bicuspidata (L.) Dum.; b136340; CB; d; new to Bridal Veil FallsCephalozia lunulifolia (Dum.) Dum.; b136341; CB; wCephaloziella divaricata (Sm.) Schiffn.; b136342; CB; wChandonanthus filiformis Steph.; b20411; WNA/EA; cChandonanthus setiformis (Ehrh.) Lindb.; b20441; CB-EA; amChiloscyphus polyanthos (L.) Corda; b136343; CB; wConocephalum conicum (L.) Lindb.; b136344; CB; wDiplophyllum albicans (L.) Dum.; b20424; CB; cDiplophyllum taxifolium (Wahlenb.) Dum.; b91021; CB; wDouinia ovata (Dicks.) Buch; b20434; WNA/ENA/WE; cFrullania bolanderi Aust.; b121766; WNA/ENA/WE/EA; scFrullania californica (Aust.) Evans; b136345; WNA; scFrullania tamarisci subsp. nisquallensis (Sull.) Hatt.; b16229; NA; cGeocalyx graveolens (Schrad.) Nees; b95635; CB; wGymnomitrion obtusum (Lindb.) Pears.; b136346; WNA/ENA/WE; wHerbertus aduncus (Dicks.) S.Gray; b136347; WNA/ENA/WE/EA; dJamesoniella autumnalis (DC.) Steph.; b136348; CB; wJungermannia atrovirens Dum.; b136349; AA; wJungermannia leiantha Grolle; b136350; NA/WE; wLepidozia reptans (L.) Dum.; b136351; CB; w125Lophocolea cuspidata (Nees) Limpr.; b136352; CB; dLophocolea heterophylla (Schrad.) Dum.; b136353; CB; wLophozia alpestris (Schleich. ex Web.) Evans; b136354; AA; am; new to Bridal Veil FallsLophozia excisa (Dicks.) Dum.; b20214; CB; wLophozia gillmanii (Aust.) Schust.; b136355; CB-EA; w; new to Bridal Veil FallsLophozia guttulata (Lindb. et H.Arnell) Evans; b80952; CB; wLophozia heterocolpos (Thed.) M.A.Howe; b79770; CB; wLophozia incisa (Schrad.) Dum.; b136356; CB; wLophozia ventricosa (Dicks.) Dum.; b136357; CB-EA; wMarsupella emarginata (Ehrh.) Dum.; b136358; CB; dMetzgeria conjugata Lindb.; b136359; CB; cMetzgeria temperata Kuwah.; b136360; NA/J; cMoerckia hibernica (Hook.) Gott.; b136361; CB-EA; cOdontoschisma denudatum (Nees ex Mart.) Dum.; b136362; WNA/ENA/WE/EA; cPeIlia neesiana (Gott.) Limpr.; b136363; CB; w; new to Bridal Veil FallsPlagiochila asplenioides (L.) Dum. sensu lato; b136364; CB; wPorella cordaeana (Hueb.) Moore; b136365; WNA/WE; dPorella navicularis (Lehm. et Lindenb.) Lindb.; b136366; WNA; cPorella roellii Steph.; b136367; WNA; cPreissia quadrata (Scop.) Nees; b136368; CB; wPtilidium californicum (Aust.) Underw.; b65568; WNA/EA; dPtilidium pulcherrimum (G.Web.) Hampe; b67657; CB; wRadula bolanderi Gott.; b18147; WNA; cRadula complanata (L.) Dum.; b102608; CB; wReboulia hemisphaerica (L.) Raddi; b136369; CB; swRiccardia chamedryfolia (With.) Grolle; b136370; CB; cScapania americana K.Muell.; b136371; WNA; cScapania bolanderi Aust.; b136372; WNA/EA; dScapania umbrosa (Schrad.) Dum.; b136373; NA/WE; dTritomaria quinquedentata (Huds.) Buch; b136374; AA; amTritomaria scitula (Tayl.) Joerg.; b86715; AA; am126Appendix COccurrence in habitats and on substrata of species collected at Bridal Veil Falls duringthe course of this study.Habitats are: ha-anthropogenic, hb-boulder slope, hc-cliffs, hf-forest, hs-streams, hp-sprayzones. Substratum are: se-vegetation, sr-rock, ss-soil, sw-wood.Species ha hb hc hf hs hp se sr ss swMossesAmblystegium serpens 1 0 0 0 0 0 0 1 0 0Amphidium californicum 0 1 0 0 0 0 0 1 0 0Amphidium lapponicum 0 1 0 0 1 0 0 1 0 0Anacolia menziesii 0 1 1 0 0 0 0 1 0 0Anomobryum filiforme 0 0 1 0 0 0 0 0 1 0Antitrichia californica 0 1 0 1 1 0 1 1 1 1Antitrichia curtipendula 0 1 0 1 0 0 1 1 0 1Atrichum selwynii 1 0 0 0 0 0 0 0 1 0Atrichum undulatum 0 1 0 1 0 0 0 1 1 0Aulocomnium androgynum 0 1 0 1 0 0 0 0 0 1Barbula convoluta 0 0 1 0 0 0 0 1 1 0Bartramia pomiformis 0 1 1 0 0 0 0 1 1 0Blindia acuta 0 0 1 0 0 0 0 1 0 0Brachythecium asperrimum 0 0 1 1 0 0 1 1 1 1Brachythecium frigidum 0 1 1 0 0 0 0 1 1 0Brachythecium plumosum 0 1 1 0 0 0 0 1 1 0Brachythecium rivulare 1 0 1 0 1 0 0 1 1 1Brachythecium rutabulum 0 0 0 1 0 0 0 0 1 0Brotherella roellii 0 0 0 0 1 0 0 0 0 1Bryoerythrophyllum recurvirostre 0 0 0 0 1 1 0 1 0 1Bryum capillare 1 1 1 1 0 0 0 1 1 0Bryum miniatum 0 0 1 0 0 0 0 0 1 0Bryum pseudotriquetrum 0 0 1 0 0 1 0 1 1 1Buxbaumia piperi 0 0 0 1 0 0 0 0 0 1Calliergonella cuspidata 1 0 0 0 0 0 0 0 1 0Campylium chrysophyllum 0 0 0 0 1 0 0 1 1 0Campylium stellatum 0 0 1 0 0 1 0 0 1 0Ceratodon purpureus 0 0 0 1 0 0 0 1 1 0Claopodium bolanderi 0 1 0 0 0 0 0 1 0 0Claopodium crispifolium 1 1 1 1 1 1 1 1 1 1Climacium dendroides 1 0 0 1 0 0 1 0 1 0Cratoneuron filicinum 0 0 1 0 0 0 0 1 1 0Dendroalsia abietina 0 1 0 1 0 0 1 1 0 0Dichodontium pellucidum 0 0 1 0 1 0 0 1 1 1Dicranella schreberiana 1 0 1 1 0 0 0 0 1 0Dicranella varia 0 0 0 0 0 1 0 0 1 0Dicranoweisia cirrata 0 1 0 1 0 0 1 0 0 1Dicranum fuscescens 0 0 0 1 0 0 1 0 0 1Dicranum scoparium 0 1 1 0 0 0 0 1 1 1Dicranum tauricum 0 1 0 1 0 0 1 1 0 1127r-I CZ>^(Z) CZ> H CZ> CZ> r-I r-I r-I CZ> CZ) CZ> CZ> CZ> CZ) CZ)^r-i H CZ> CZ> CZ) CZ>^CZ> CZ> T-4 r-I H <Z> r-4 r-i^r-4 r-4 (7) r-I r-1 CZ) CZ> r-I r-I CZ) T-4 r-I CZ>r-4 r-I^1-4 r-I H CZ> r-1 r-I r-I CZ) CD H r-A^(7) r-I CZ> CZ> r-4^CZ) r-I H r-I CZ> H^r-I T-4 CZ> r-I H CZ> H T--1 <Z> T-4 CD CD CD CD CZ> r-I r-4 r-4 r-I r-Ir-I %-1 T-IHT-Ir-lr-4 CZ) CZ> %-1 (7) H r-4 r-I CZ> r-i^r-i CZ> r-I r-i H r-I CZ> Cf) r-I r-I CZ> T-1 r-I r-4 CZ> r-I r-i CZ> CZ) CZ> CZ> CZ> CZ) CZ> T---1 CZ) CZ) r-I r--ICZ) CZ) CZ> CZ) CZ> CZ> CZ> <Z> T-I r-4 CZ>^T-4 r-I CZ) CZ> CZ> CZ) r-i r-i r-I^r-I CZ) CZ> r-i CZ>^H r-I CZ> r-I CZ> T-I t-1 r-i CZ> 1-1^r-I r-i CZ) CZ) r-A H r-ICZ) <Z> r-i r-I CZ) H CZ) CZ> CZ) <Z> 1-4 CZ> r-I CZ> CZ> CZ) CZ) <Z> CZ) CZ> CZ) H CZ> H <Z) CZ) CZ> CZ> CZ) CZ> H r-I r-I CZ>^CZ> CZ) CZ> CZ> CD CD CD CD r-I CZ> CZ> 1---1 CZ)I-4 CZ> CZ) 1-.1 CZ> CZ) CZ> CZ> CZ) T-4 CZ> <Z>^r-4 CZ> CZ> CZ>^r-I r-4 CZ> r-I CZ) CZ> CZ> CZ> CZ>^r-I T-1 CZ> CZ> r-I CZ) r-I CZ> CZ> CZ> CZ> CZ> <Z) <Z> CZ> r-1 CZ> CZ> r-I CZ)<Z> CZ> (7) CZ> CZ) CD CZ) r-I r-I^CZ> T--1 r-I CZ> CZ> H (7) CZ> H T-A r-4^r-i CZ> r-I H CZ) r-A r-1 1-1 r-4 r-4 r-I T-4 r-I r-i r-A r-4^r-i r-i r-I H CZ> CZ) r-4 r-I HC:) H r-I H r-I r-i r-I CZ> CZ> CZ> CZ) (-.1 CZ) r-4 CZ> r-i r-A^CZ> r-i^r-4 H r-I r-I CZ> CZ> H r-4 r-i r-4 CZ> T-I CZ> H r-A CZ> CZ> CZ> CZ> CZ) CZ> CZ> r-1 r-I CZ> r4 CZ)r-I <Z> CZ) r-4^CD CZ) CZ> r-I r-4 CZ> <Z> CD CZ> CZ> r-I r-I 1--1 CZ) r-I r-4^r-i CZ> CZ> CZ> CD r-4 r-I CZ) r-I CZ> CZ> CZ> CZ> 1-4 CZ> CZ> <7) CZ) CZ> <Z> CZ) CZ) CZ>^r-I r-ICZ) CZ> r-I CZ) CZ) CZ) CZ> CZ> <Z> T-4 CZ> CZ) T-4 CZ>^CZ) CD CZ> CZ> r-i CZ> CZ) CZ> CZ) CD CZ> r-I (Z) CD CZ> CZ) CZ> CZ> CZ> <Z) CZ> CZ> CD CD CZ> CD CD CD CD CZ) r-4 r-I CZ>a)4-)g^U)^U)^0^10- 1-1- H -0 co^5^9^u)^5 -,-.1 u) (T).,-1^rn -H a) r...) -H^a) -H^a) u)^-1-1^a)^5^(13U)4^g ts)-H .-1^z co^o^9:, (1) ■-I^(/)^ro a) -H^g —1^r-1^g 4..)7^1-1 t-A (1)^Rs^5 gU)^u)^.., -H P^-H .-I^00^0 0 -H a) (15^m $-1^-.-1^(I)^S-1 g - H CD 0 -I-> 4-)^(11 -L>U)   H^03^Z 0 U) (I) U) $-I - H^- H 0 0 t:11 -I-)^(I) - rl^U100a)^a) -H ■-I ,0 -I-1 (C) 0 -H^rti 4..)^,-1 -H _i_) rt5 ni C.) 43.) ni f—i W 4 W 4-1^0) 0 4 1--1 -1--) rt:5 75 u) 0^-H 0 S-I ,-0 -H^-H^tfl U) r--I C.) c...) -H^-H t)) $4 u)^U)— WO ozi a 05 a) .L) o a) rt5 54 ,--1 -H rci a) .L.) $4^E (4-1 z S-1 W -H S-I 1-1 b)^U)^0 -H u) (i r.-1 o >1^104 4-) 0 a) u) u) Zrc, r-A -r1 a) pa -H c) (a, 4.) ,-1 ..1 0 :i C) 4^n3 Q)^0^f>4 ;i ,--i 0 .0 4 () NI 01 CD gl () 9Z1 -r4 4-i o5 0 nj 0^;J -1-1 4 Z (D CD -r-I (D 7 .0 t4^Ei Ei^-1-1^-r1 gl a) r-4 a) 4^:: irrii -: ::0 —I^(5 -1 0 0 it CD g _i) u)^rt50-H^- rti U ,--i 04^u) U -H 4 0 W- i—i b) - H i—i 0 rti^0 Cli 4 C)i -.,s 2 .10 g 0 g ..8 g g ,--1 (19, g g •r.4 44) pi >•1 coo E g 2 roiS —10.1 2 5 9 5 5 5 5 034-.) 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This table provides synonyms to the names used in Lawton or Godfrey.Names used in this report are to the lefthand side, synonyms used by Lawton or Godfrey areto the righthand side. In the case of species not treated by these two authors, alternatereferences are provided.Anomobryum filiforme = Pohlia filiformisBarbula amplexifolia - not treated in Lawton (1971), see Zanders (1979)Barbula convoluta var. gallinula - not treated in Lawton (1971), see Zanders (1979)Bryoerythrophyllum recurvirostre = Didymodon recurvirostrisCampylium polygamum = Amblystegium polygamumDidymodon fallax var. reflexus = Barbula reflexaDidymodon rigidulus var. gracilis = Barbula acutaDidymodon vinealis var. flaccidus = Barbula vinealis var. flaccidaDitrichum crispatissimum - referable to D. flexicaule, see Frisvoll (1985) for detailsEncalypta rhaptocarpa = E. vulgaris var. rhabdocarpaHerberta adunca = Herbertus aduncusHymenostylium insigne - not treated in Lawton (1971), see Zanders and Eckel (1982)Hymenostylium recurvirostre = Gymnostomum recurvirostreIsopterygiopsis pulchella = Isopterygium pulchellumIsothecium myosuroides = I. stoloniferumLeucolepis acanthoneuron = L. menziesiiMetaneckera menziesii = Neckera menziesiiMnium ambiguum = M. lycopodioidesMnium thomsonii = M. orthorrhynchumPalustriella commutata = Cratoneuron commutatumPlagiopus oederiana = P. oederi131Plagiothecium cavtfolium = P. roeseanumPlatyhypnidium riparioides = Eurhynchium riparioidesPolytrichastrum alpinum = Polytrichum alpinumPorotrichum bigelovii = Porothamnium bigeloviiPorotrichum vancouveriense = Bestia vancouveriensisPseudoleskea stenophylla = Lescuraea stenophyllaPseudotaxiphyllum elegans = Isopterygium elegansRacomitrium spp. = Rhacomitrium spp.Racomitrium canescens s.str. - not treated in Lawton (1971), see Frisvoll (1983), or Hill(1984)Racomitrium elongatum - not treated in Lawton (1971), see Frisvoll (1983), or Hill (1984)Racomitrium lawtonae - see discussion under R. heterosti chum var. heterostichum in Lawton(1971), for more information see Frisvoll (1988)Racomitrium pacificum - not treated in Lawton (1971), see Frisvoll (1988)Sanionia uncinata = Drepanocladus uncinatusSphagnum spp. - not treated in Lawton (1971), see Ireland (1982)Schistidium apocarpum = Grimmia apocarpaTrichodon cylindricus = Ditrichum cylindricumUlota obtusiuscula = U. crispa var. alaskanaZygodon viridisssimus = Z vulgaris132Appendix ESimilarity between habitat/substratum combinations based on composition of theirrespective moss flora using the Jaccard Index of Similarity.Habitat substratum combinations are named with two letters. The first letter refers to thehabitat, where a=anthropogenic, b=boulder slope, c=cliff, f=forest, s=stream, and p=sprayzone, the second letter refers to the substratum, where e=epiphyte, r=rock, s=soil, andw=wood.Habitat/^ Jaccard index of similaritysubstratum (X 102)combinations'as be br bs bw ce cr cs fe fr fs fw se Sr ss sw pe pr ps pwar (7 )^10 17 6 0 0 0 10 3 4 11 8 5 20 21 7 10 0 0 0 5as (15) 0 7 9 5 0 7 10 11 16 23 9 11 7 15 15 6 10 0 4be (1 ) 2 0 0 0 2 0 2 0 4 3 20 6 0 6 0 0 0 0br (54)^ 18 6 2 24 12 23 7 15 15 6 11 6 11 2 4 2 5bs (30) 12 0 10 6 14 6 19 13 0 6 7 3 0 4 4 0bw (13) 02 0 10 0 3 800  0 0 0 005ce (1 )^ 2 3 0 7 0 0 0 6 0 0 0 0 0 8cr (61) 42 17 13 9 7 6 19 8 14 4 10 0 0cs (43) 6 7 8 3 6 12 12 12 0 12 12 23fe (60)^ 9 23 50 9 7 8 19 2 6 2 9fr (14) 19 7 6 11 10 11 6 5 5 8fs (24) 25 12 8 15 22 4 11 0 3fw (45)^ 6 7 820 3 3 0 5se (5 ) 17 18 17 0 8 0 6sr (18) 26 19 0 9 5 4ss (12)^ 26 0 14 7 11sw (18) 0 20 5 21pe (3 ) 000pr (9 )^ 0 24ps (8 ) 11pw (21)s Numbers in brackets represent total number of moss species found ineach habitat substratum combination.Note: Habitats ae, aw, and cw had no species and were dropped fromthe analysis.133Appendix FA note of caution.The following species are documented by specimens at the herbarium of theUniversity of British Columbia and are labelled from Bridal Veil Falls. Based on the habitatsin the area, the thoroughness of the field studies of the past two seasons, and the ecology ofthese species, it seems very possible that the specimens were not collected from Bridal VeilFalls. It is probable that they are from some site closer to the coast, probably around HoweSound, where the collector (J.P. van Velzen) also made collections. All of the species notedare known from the latter area and are unknown (except for van Velzen's specimens) fromBridal Veil Falls. It is not impossible that the specimens are correctly labelled, but on thebasis of field experience at Bridal Veil Falls, their presence there should be considered asuncertain until new collections are made.These species are:^Andreaea rupestris Hedw.;Chandonanthus filiformis;Chandonanthus setiformis;Diplophyllum albicans;Douinia ovata.Unlike the four other species, Andreaea rupestris had not been included in the florabecause the herbarium material was not available for examination. However, B.M. Murray(University of Alaska Museum) recently confirmed that van Velzen's records of A. rupestrisfrom Bridal Veil Falls (UBC B20440, B20431, B20421) were correctly identified.A. rupestris is easily distinguishable from the other species of mosses found at BridalVeil Falls. It is a small plant, brown or reddish-brown in colour, with ecostate leaves, and asporophyte which opens along four longitudinal slits.134


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