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The phytogeography and ecology of the mosses within the San Juan Islands, Washington State Harpel, Judith Strachen 1997

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THE PHYTOGEOGRAPHY A N D ECOLOGY OF T H E M O S S E S W I T H I N T H E S A N J U A N I S L A N D S , WASHINGTON  STATE  by JUDITH S T R A C H E N H A R P E L B . S . California State Polytechnic University, Pomona, 1974 M . S . California State Polytechnic University, Pomona, 1980  A THESIS S U B M I T T E D IN P A R T I A L F U L F I L L M E N T OF T H E REQUIREMENTS FOR T H E D E G R E E OF D O C T O R OF P H I L O S O P H Y in T H E F A C U L T Y OF G R A D U A T E STUDIES Department o f Botany  We accept this thesis as conforming to the required standard  T H E UNIVERSITY OF BRITISH C O L U M B I A  A p r i l 1997 © Judith Strachen Harpel  1997  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department  or  by  his  or  her  representatives.  It  is  understood  that  copying  or  publication of this thesis for financial gain shall not be allowed without my written permission.  Department The University of British Columbia Vancouver, Canada  DE-6 (2/88)  Abstract Floristic work on bryophytes i n the state o f Washington has been confined mostly to the mainland with little information available for the San Juan Islands. After four years o f field work and an extensive search o f historical records from herbaria throughout the region, the San Juan Islands prove to contain a diverse moss flora within a small geographic area; this flora consists o f 224 species and varieties, 33 families and 97 genera. Four species Drepanocladus crassicostatus, Orthotrichum hallii, Tortula papillosa  and Tortula laevipila var. meridionalis are reported new for the State o f  Washington. Tortula laevipila var. meridonalis is new for the United States and represents the second North American location. Detailed ecological observations were made for each collection and distributions for each species have been mapped. Keys are presented to both genus and species. The bulk o f this flora is composed o f circumboreal species that are derived from a once widespread Arcto-Tertiary flora. During the Pleistocene these islands were completely glaciated and the present flora represents, therefore, species that have migrated back into the region predominantly from southern refugial sites probably during the Hypsithermal Interval described by Deevey and Flint (1957). A cooling trend about 2000 yr. B . P . probably caused the southern element species to retreat southward throughout the region with fragments persisting only in those areas where favorable conditions also remained. The San Juan and adjacent islands can be interpreted as a "modern" refugium for southern mediterranean type climate species.  ii  Table o f Contents Abstract  ii  Table of Contents  iii  List of Tables  v  List of Figures  vi  Acknowledgment  v i i i  Forward  x  Chapter One Introduction  .,..1  Chapter Two Early bryological work within the San Juan Islands Chapter Three Description o f the study area Climate Geology and soils Vegetation Late Pleistocene and Holocene vegetation and climate  3 15 15 21 33 35  Chapter Four Methods and Materials  39  Chapter Five Ecology  48  Moss species as related to habitat and sub-habitat  48  Moss species as related to substratum Moss species as related to geologic unit Unusual habitats and substrata Limestone deposits Bogs and bog like habitats  59 68 72 72 73  Chapter Six Phytogeography  79  Floristic history Floristic patterns World continuous distributions Disjunct world distributions Pacific Northwest distributions  81 82 85 91 98  Chapter Seven Taxonomic Considerations  115  Preparation o f keys Problematic taxa  115 116 iii  Historical taxa Taxa new to Washington K e y to genera Keys to species Chapter Eight Summary and conclusions  119 119 121 134 ;  153  Literature Cited  157  Appendix A  168  Appendix B  169  Appendix C  170  Appendix D  175  Appendix E  180  Appendix F  185  Appendix G  190  Appendix H  195  Appendix 1  200  Appendix J  205  Appendix K  211  Appendix L  216  Appendix M  217  Appendix N  218  iv  List o f Tables Table 1. Temperatures and precipitation from known states in the San Juan Islands  19  Table 2. Geologic formations in the San Juan Islands  24  Table 3. Glacial history in the Puget Trough  27  Table 4. Soil types found in the San Juan Islands  30  Table 5. Soil units in the San Juan Islands  31  Table 6. Pollen records in the Puget Trough  37  Table 7. Collecting locations  .  r  40  Table 8. Number o f sections visited by sub-habitat on each island  46  Table 9. Habitats and sub-habitats for the San Juan Islands  49  Table 10. Islands and sub-habitats from which collections were taken  50  Table 11. Moss diversity in the habitats  52  Table 12. Definitions o f substrata  60  Table 13. Moss diversity according to substratum  61  Table 14. Moss diversity as related to geological unit  69  Table 15. The world distributions o f the mosses in the San Juan Islands  80  Table 16. The Pacific Northwest distributions of the mosses in the San Juan Islands  80  Table 17. W o r l d phytogeographic elements in the Northern Hemisphere  86  Table 18. Pacific Northwest phytogeographic elements  86  Table 19. The relationship between the number of mediterraean climate species found in each habitat 107 Table 20. San Juan moss diversity comparisons to other areas  Ill  List of Figures Figure 1. Location map for the San Juan Islands  16  Figure 2. Isohyetal map o f the San Juan Islands..  18  Figure 3. Geology o f the San Juan Islands  23  Figure 4. Marine limit at about 13,500 yr. B . P . , submerged area is shaded  28  Figure 5. Soils in the San Juan Islands  29  Figure 6. M a p o f collecting locations in the San Juan Islands Figure 7. Pacific North American Region  , .,  44 84  Figure 8. The world distribution o f Hylocomium splendens  87  Figure 9. The world distribution o f Grimmia laevigata  87  Figure 10. The world distribution of Aulacomnium palustre  89  Figure 11. The world distribution o f Hypnum dieckii  89  Figure 12. The world distribution oiDendroalsia  92  abietina  Figure 13. The world distribution o f Andreaea megistospora  92  Figure 14. The world distribution o f Orthotrichum hallii  94  Figure 15. The world distribution o f Heterocladium  macounii  94  Figure 16. The world distribution o f Plagiothecium  denticulatum  97  Figure 17. The world distribution of Antitrichia  californica  97  Figure 18a. The Pacific Northwest distribution o f Pleurozium schreberi  100  Figure 18b. The Pacific Northwest distribution o f Ulota phyllantha  100  Figure 19a. The Pacific Northwest distribution o f Claopodium bolanderi  102  Figure 19b. The Pacific Northwest distribution o f Trichostomopsis  102  australasiae  Figure 20a. The Pacific Northwest distribution o f Orthotrichum hallii  104  Figure 20b. The Pacific Northwest distribution of Alsia californica  104  vi  Figure 21. The distribution of Drepanocladus and Juniperus scopulorum  crassicostatus,  Orthotrichum  Figure 22. The distribution o f Dendroalsia abietina and Quercus garryana  vii  hallii 105 109  Acknowledgments I would like to thank Dr. W i l f Schofield for his assistance and guidance throughout this project and for his tireless enthusiasm for fieldwork. Thanks are also extended to my committee members, Drs. Gary Bradfield, Gilbert Hughes and Gerald Straley for their support and help during this research. Field work was partially supported by a grant to Dr. Schofield from the National Sciences and Engineering Research Council o f Canada. Drs. Jan Janssens and R o n Pursell provided critical assistance in determining troublesome taxa. Special thanks goes to Rene Belland and Olivia Lee for being "sounding boards" during the identification process. Suggestions for collecting locations were provided by Dr. Eugene K o z l o f f and the staff at the Friday Harbor Biological Laboratory. Computer "crisis counseling" was provided by Penelope (Lebby) Balakshin in the U . B . C . Botany Department. . Sarah Gage gave invaluable assistance and access to the bryophyte collection at the University o f Washington Herbarium. Transportation and collecting permits were provided by the United States Fish and Wildlife Service, United States National Park Service, United States Bureau o f Land Management, Washington State Parks, The Nature Conservancy and The San Juan Preservation Land Trust. Numerous private property owners graciously allowed us access to their land. Ruthie Johns and her family provided housing, transportation and excellent contacts with people throughout the islands. Grateful thanks are extended to all of these individuals and organizations. I would also like to thank my son Chris for helping with the fieldwork, and sharing his geological background which provided additional insight into my research.  viii  Finally a special thanks goes to my parents, who always encouraged me to dream as a child, and to my husband whose unfailing encouragement and support both in the field and during the long periods while I was away from home allowed me to fulfill those dreams.  ix  X  CHAPTER 1 Introduction Floristic work on bryophytes in the State o f Washington has been confined mostly to the mainland, with little published information available for the San Juan Islands and no attempt has been made to synthesize the available information. Earlier researchers in the state have included: T . C . Frye, who worked on the hepatics, Sphagnum, Racomitrium,  and the  Polytrichaceae o f the Pacific Northwest, E . Lawton, who published a Moss Flora for the Pacific Northwest (Lawton 1971), and established a set o f keys for the moss flora, but provided a very general picture o f the distribution within the state. It is apparent that some areas in the state have not been extensively collected. R . R . Ireland collected in the state while he attended the University o f Washington, but he has published only a portion o f his data. G . Jones, W . Suksdorf, and C . Piper were early botanists who collected bryophytes but are better known for their vascular plant work. Mrs. Betty Higinbotham, a bryologist from Washington State University, Pullman, spent her summers on San Juan Island, where she made some collections and has published a short list of the more common mosses in a local publication (Higinbotham 1985). Finally W . B . Schofield has collected within the adjacent G u l f Islands and on Vancouver Island, but has never collected extensively in the San Juans. Thus a data base exists for the G u l f Islands and the southern part of Vancouver Island but not for the San Juans. The lack o f bryophyte floristic work combined with the unique climate, geological and vascular plant history o f the San Juan Islands identified this area as one with promising research possibilities. Besides the lack o f prior work and the unusual climatic and physical features that the San Juan Islands offer, there are advantages and disadvantages to working on islands. The primary advantage to developing and interpreting a flora from a group o f islands is that the boundaries o f the flora are clearly defined, and when there has been little or no disturbance to the island it presents an excellent opportunity to study natural floristic patterns. Disturbance and accessibility are the most significant disadvantages to working in a group o f islands. Both human and animal disturbance w i l l have a greater impact on small areas and this 1  may result in a change o f species composition. Lack o f access can prevent species from being found. The present study was designed with the following primary objectives. 1. To develop a complete moss flora for the San Juan Islands through field work and a careful search o f historical records. 2. To collect data on the habitat, substrata, and geological unit that each moss was found in or on, and to compare these relationships. 3. To determine the geographical distributions o f the species and discuss their possible origins. 4. To prepare keys to the genera and species comprising the flora. 5. To map the distributions within the islands for each species This study was designed to develop a comprehensive data base that can be used to assess any future changes that the islands may undergo. A l s o when equally comprehensive data exist for the adjacent areas, better conclusions can be made concerning the significance o f the flora of these islands in relationship to these areas.  2  CHAPTER 2 Early Bryological Work Within The San Juan Islands  The first collections o f plants were made i n the Pacific Northwest approximately two hundred years ago. The two naturalists who first botanized the area were Tadeo Haenke and Jose Mocino. (Piper 1906.)  Haenke, an Austrian, was with the Malaspina Expedition o f  1791 and collected both flora and fauna; it is not known whether he obtained any bryophytes during his travels. Mocino, a botanist with the 1792 Royal Expedition o f N e w Spain lists six bryophytes in the catalogue o f animals and plants section o f his Noticias De Nutka (Mozino 1970). Although he collected a few bryophytes, all o f his work was confined to the Nootka Sound area o f Vancouver Island and he apparently never visited the San Juan Islands. While the Spanish played a significant role in the exploration and naming o f these islands, they contributed no bryological information. Archibald Menzies was probably one o f the most important botanists who worked in the region during this time. He was with Vancouver's 1792 Expedition and is best known for his extensive vascular plant collections. During his visit to the area, according to his journals, he collected some vascular plants from several o f the islands (Newcomb 1923). He also lists the bryophytes that he collected during the expedition. To determine the specific localities o f these bryophytes, a request was made to the Royal Botanic Gardens in Edinburgh (where the collections are held) for label information, but the results were disappointing. Most o f his bryophyte collection labels were either "Northwest Coast o f America or WestCoast o f North America" (Long in litt. 1994). It is therefore, difficult to determine whether Menzies actually collected bryophytes from the San Juan Islands although he certainly visited them. Although both John Scouler and David Douglas were in the Pacific Northwest in 1825 with the Hudson Bay Company, and collected bryophytes, neither o f them visited the islands within San Juan County. When Scouler left the mouth o f the Columbia River to head  3  north, Douglas remained behind to explore that area. According to Scouler's journals he was on both Cypress and L u m m i Islands in August o f 1825 (Young 1905), but there is no record that he collected any bryophytes or visited any o f the other local islands. Therefore it is reasonable to assume that neither o f these botanists collected bryophytes within San Juan County. The next major research event was the visit to the area by the United States Exploring Expedition led by Captain Charles Wilkes in 1841. This expedition is often referred to as the Wilkes Expedition; it was charged by Congress to explore all o f the Southern Ocean. Ultimately the expedition proceeded up along the west coast o f California and what was then called the Oregon Territory. According to Jackson (1985) the primary focus o f this expedition was to survey and map all o f the lands within the Southern Ocean and, i f time permitted, to allow the civilian scientists to carry on their research. The ships under Wilkes' command were dispatched to various parts o f the region at the same time; therefore, the scientists were not always able to explore all o f the areas being mapped. Keeping this in mind when reading his report, it is possible to determine how briefly Wilkes visited the San Juan Islands. After crossing the Straits o f Juan de Fuca with a party o f seven small boats, Wilkes and his men set out to explore: "On the 26th (July) we began the survey o f this labyrinth of islands, which was continued the next day," (Wilkes 1849). They finished the survey on the 28th and when the news o f the shipwrecked Porpoise arrived, they cut short the rest o f the survey. Wilkes (1849) states: "I regretted that I had been deprived o f the opportunity o f examining the southeast end o f Vancouver Island which I have reason to believe offers many fine harbors. Three days more would have enabled me to accomplish this portion to my satisfaction". During his entire narrative o f this survey, Wilkes made no reference to stopping on any o f the islands for research purposes, or that any o f the civilian scientists who were with him did so. The two naturalists assigned to this expedition, M r . Pickering and M r . Breckenridge collected both vascular plants and bryophytes, yet it appears that they did not collect within 4  the islands. According to Sayre (1975a) all o f the bryophytes from the entire expedition were sent to W . S . Sullivant for determination. A review o f Sullivant's work, published in 1859 by A s a Gray, showed that the closest collection locality for all o f the material was listed as Puget Sound, Oregon (Tan in litt. 1994). This locality probably was the Fort Nisqually area, which is now in Washington State. Wilkes, like the Spanish explorers before him, was also responsible for naming many o f the islands. However, his expedition did not contribute significantly to the bryological information for the region. In 1846 a treaty between the United States and Great Britain established the boundary between the United States and Canada as the Forty-Ninth Parallel o f North Latitude. This boundary extended westward to "the middle o f the channel which separates the continent from Vancouver's Island, and thence southerly through the middle o f said channel and Fuca's Straits, to the Pacific Ocean" (Klotz 1917). While debate over who held jurisdiction over the San Juan Islands continued until 1872, these islands were included i n the joint British and United States Forty-Ninth Parallel Boundary Survey carried out between 1857 and 1861. The primary focus o f this survey was to clearly define the boundary from the Strait of Georgia eastward to the Cascade Mountains. Dr. David L y a l l , a Scottish physician and naturalist was assigned to this survey as ship's surgeon and botanist. W h i l e L y a l l collected mainly vascular plants, his interest in cryptogams is revealed in a paper read to the Linnean Society o f London on June 18th 1863. In this paper he states, "I have been fortunate in securing the services o f M r . W i l l i a m Mitten, A . L . S . in the arrangement and naming o f my extensive collections o f Mosses, Hepaticae, and Lichens, which thus have the value o f being named by one o f the most able and assiduous Cryptogamists in England" (Lyall 1864). During the summers o f 1858 and 1859 Lyall spent part o f his time while on board the Plumper, surveying the area around the southeast corner o f Vancouver Island and among the surrounding islands. It was during one o f the numerous stops that L y a l l collected a liverwort from Orcas Island. Mitten (1864) in his paper on the "Bryologia o f the Survey" identifies Lyall's liverwort as Frullania  tamarisci L . and gives the location as Vancouver and Orcas 5  Islands. Although Sayre (1975b) states that an intact set o f Lyall's bryophytes from the entire survey consisted o f 116 mosses and 19 liverworts, Mitten's paper (1864) mentions only the one San Juan Islands site. Therefore it appears that Lyall did not collect any mosses from these islands. H i s liverwort record, on the other hand, represents the earliest documented bryophyte record from these islands. Benson (1986) states that "before the 1880's neither individuals nor local institutions pursued natural history work in the Puget Sound region. But by the early 1900's organized studies in the natural sciences were conducted through the zoology, botany and geology departments at the University o f Washington, the marine station o f the University at Friday Harbor and the new Washington State Museum", Rigg (1929) also points out that most o f the botanical field work done after 1860 in the state was carried out by local residents and they appear to have worked mostly on the mainland. Therefore it is not surprising that for the next thirty-four years after Lyall's work there was little or no botanical research activity within the San Juan Islands. It was not until 1892, when Louis F. Henderson spent several days in July collecting around the Eastsound and M t . Constitution areas o f Orcas Island (Sarah Gage, pers. comm. 1994), that botanical research occurred within the islands. H i s primary interest at this time was to obtain vascular plants for the upcoming World's Fair. Although he did collect bryophytes (Vitt et al. 1985), it would be difficult to determine i f any were collected from Orcas Island during his visit because most o f his original collections prior to 1906 were lost in a fire at the University o f Idaho (Piper 1906). The advent o f the late 19th Century and the early 20th Century brought about change to both natural history and the San Juan Islands. American natural history during the 19th Century was dominated by amateurs and was a very popular pastime (Benson 1986). Burnham (1971) further comments that "local natural history societies flourished everywhere, even near the frontier, to foster and express the scientific interests o f well educated and even self-educated Americans". 6  It was in this atmosphere that the Young Naturalists Society was formed in 1879 by a group of young Seattle area men. This group by 1885 had developed close ties with the University o f Washington and was rapidly assembling substantial field collections. In 1894 new members included school teachers, several physicians and Trevor Kincaid (Benson 1986). When this society disbanded in 1904 its collections formed the beginnings o f what is now the Burk Museum, and it had provided the ground work for the development o f the Friday Harbor Biological Laboratory under Kincaid's guidance. Hayner (1929) defines the period o f development from 1881 to 1910 as the "Village Stage" for the San Juan Islands. During this stage, the population increased and several major industries were established. This increase in both population and economy brought about the construction o f new roads, and easier access to and from the mainland. Thus the "Bryological Era" had begun within the San Juan Islands. The "Bryological Era" in the San Juan's can be divided into two distinct periods based upon historical collecting information obtained from the herbaria o f the University o f Washington, Western Washington University, Portland State University, and the University of British Columbia. The first period starts in 1902 and continues until 1908. In 1902 and 1903 all o f the collecting was done by either Dr. John W . Bailey or M r . A . S . Foster. Dr. Bailey was a well known Seattle area physician who was extremely interested in bryophytes. He contributed to both A . J . Grout and J . M . Holzinger's moss Exsiccata and served as Secretary of the Sullivant Moss Society for several years (Frye, 1933). There is some confusion as to the identity o f M r . A . S . Foster; Steere (1978) lists him as Adriance Sherwood Foster 1901- 1973, but this is impossible, as herbarium material collected from the San Juan's by A . S . Foster starts in 1902. While Foster always used his initials when he prepared his labels and published articles, there is some indication that the A . S . initials stood for Albert Scott. Holzinger (1921) listed Foster as a "teacher in the public schools o f the state of Washington....". When a request was made to the Washington State Superintendents o f Public Instructions Office, they reported that their records for the era 7  listed a teacher certificate number for an Albert Scott Foster. Unfortunately all o f the early information was sent to the state Archives Office and his file was disposed of. The only record that they have for a Foster was found in the Register o f Life Diplomas Granted by the State o f Washington in 1902. He is listed as A . S . Foster, 48 years old, and from the state of Indiana, at the time he was given his life teaching certificate. In one o f his papers Foster makes reference to "my boyhood days in southern Indiana ..." (Foster 1904), thus confirming that he was the same teacher who was interested in bryophytes. While Foster published papers on both vascular plants and bryophytes, his interest in bryophytes is evident from his vast exchange o f material with the well known bryologists John Holzinger and A . J . Grout. Foster sent material from Waldron island to Grout for determination and Grout used Scleropodium colpophyllum as number 392 in his exsiccata o f North American M u s c i Pleurocarpi which was issued, according to Sayre (1971), in June o f 1912. Foster's contributions to northwestern United States bryology are also noted in a paper on the early history o f the Sullivant Moss Society by Annie M o r r i l l Smith (1917), "Then in field work we might fairly claim to have promoted exploration when we recall the collections o f A . S . Foster in Oregon and Washington  During his work in the San Juan  Islands, Foster collected on the following islands: San Juan, Henry, Orcas, Waldron, and he was the only person ever to collect bryophytes on Stuart. The contributions that both Bailey and Foster made to Northwest bryology also can be documented by Holzinger's proposing the new name Bryum baileyi for a species o f Bryum collected by Bailey in the Seattle area. Holzinger (1905), states "I propose ... in recognition o f my very efficient bryological friend Dr. John W . Bailey o f Seattle...". Later that same year Holzinger changed the name to Bryum fosteri, as the name B. baileyi had already been used. Other collectors in this region during this early period included Mrs. Huggens, M r s . Thomas, A . S . Pope, and J.B.Flett, but it was primarily Foster and Bailey who were responsible for most o f the early collections. It is interesting to note that the early pre-1904 collecting activity corresponds neatly to when the Young Naturalists Society was still active. Although there is no evidence that 8  Foster, Bailey or any o f the other early collectors were members o f the Society, their interests fit in well with the current hobby atmosphere o f that era, and it is possible that they were members. B y the time that the Society disbanded in 1904, the University o f Washington had opened the Puget Sound Marine Station in Friday Harbor on San Juan Island. Even though the idea o f a Marine Station was established in 1902, it was not until 1904 that the first classes were taught by Trevor Kincaid and Theodore C . Frye. (Howard 1963). Although formal classes taught on a regular basis did not occur until the summer o f 1906 (Benson per comm. 1994), the influence o f the Station began in 1904 when Frye started collecting bryophytes from the islands. Since early records for the Station are relatively scarce it is difficult to determine i f formal bryophyte classes were offered before 1923, but based on the herbarium data it seems very unlikely. The last major bryophyte collecting activity during this first period occurred in 1908, after which little collecting was done until 1919. During this period o f bryological inactivity, Frye collected briefly i n Roche Harbor, San Juan Island on July 11th 1911 and again in July o f 1915 with Goete W . Turesson (the director o f Algae at the Station) on M t . Constitution, Orcas Island. These two days represent the only bryological field work done within these islands between 1910 and 1918, according to the herbarium records. This relative lack o f collecting may be the result o f Frye's absence from the Station as well as the negative influence o f World War I. In 1913 Frye and G . B . Rigg led a survey o f the Alaskan kelp beds to look for a new source o f potash (Howard 1963). It was also during this time that A . S . Foster was asked to j o i n the expedition and, when time permitted, they collected Alaskan bryophytes (Holzinger and Frye 1921). Theodore C . Frye was by far the most significant collector o f bryophytes in the San Juan Islands. It is interesting to note that the two periods o f bryological activity correlate with his being director o f the Station from 1905 to 1909, and again from 1914 to 1930. Although, according to herbarium data, Frye spent 91 days collecting within the islands he usually went back to the same locations and therefore did not collect extensively throughout  9  the area. There are also problems with interpreting some o f his labels and, combined with the lack o f good field journals, it is often difficult to relocate some o f his old collecting sites. In spite o f these difficulties, Frye's enthusiasm for plant ecology, bryology and teaching is reflected in the second period o f the "Bryological Era" which began in 1919 and continued until 1930. From 1919 until 1930 all o f the bryological research was directly related to the Station. Florence Spaulding, a student who took the Botany Research Class at the Station in 1920, began collecting bryophytes on Blakely Island i n July o f 1919. While there, she collected Eurhynchium oreganum and Fontinalis antipyretica.  The north side of  Shaw Island was visited on July 7th 1921 by Anna M . Daugherty, also a student at the Station in 1920. She collected the following species o f mosses: Anacolia menziesii,  Bartramia  pomiformis, Neckera douglasii, Metaneckera menziesii, and Ulotaphyllantha.  Their  independent visits appear to be the only early bryophyte records for both o f these islands. In 1922, during the month o f July, Frye collected i n the Friday Harbor area and Daugherty visited the M t . Constitution area on Orcas Island. In 1923, the herbarium records indicate that a formal course in bryophytes was offered. During this time Frye was the instructor, and Daugherty (now a high school teacher in Seattle) was the teaching assistant. This class spent 34 days in the field and collected on Orcas, San Juan and Sucia Islands. Herbarium labels list the following people as collectors: Lois Clark from the University o f Idaho, A . M . Daugherty, T . C . Frye, Lena Hartage, a University o f Washington student, C M . Roberts from Pennsylvania State University, M . Wentworth, a student and Station Librarian, Catherine Smith, an Auburn H i g h School teacher, and Jean Berger, a Bellingham H i g h School teacher. Many o f the labels provided only "Class o f 1923". Dr. Bailey also collected in 1923, but it is unlikely that he was enrolled in the class. This also was the year that the University began work on the new buildings at the Point Caution location, which ultimately provided more laboratory and classroom space. Relatively little is known about the bryophyte collections o f 1924. A l l o f the herbarium labels list either Frye, Hartage or the "Class o f 1924" and represent collections 10  from only San Juan and Orcas Islands. They spent a total o f 10 days in the field and it is doubtful that a formal bryophyte class was held that year. The bryophyte class o f 1925 provides the most historical information about itself. According to an unpublished student report in the University o f Washington's Department of Biomedical History Archives, this class was "notable for the number o f new species which were collected in the vicinity o f the station and on the islands". Dr. Frye was listed as the Director o f Bryophytes, Dr. Lois Clark taught the liverworts and Chas. M . Roberts served as the teaching assistant. O f the six students, all but one o f them were either Junior H i g h or High School Biology or General Science teachers. These students were as follows: H . A . M i l l i c a n from Y a k i m a H i g h School, Sylvia E . Smith from Roosevelt Junior H i g h in Decatur Illinois, Estella Peterson from Shelton H i g h School, Glen Osborn from Bellevue H i g h School, Doris M u l l e n from Wenatchee H i g h School and finally Carra E . Horsfall from Reed College in Portland Oregon. Doris M u l l e n appears to have been a very discriminating collector and she is responsible for several historical collections from San Juan Island, including Andreaea rupestris and Climacium dendroides. Later she was the teaching assistant in the Normal Botany and Algae Classes taught in 1929 and 1940. This class also apparently traveled more throughout the islands and collected on Orcas, San Juan, Spieden and Sucia. In 1926 the only bryophyte collections were made in July by Frye and J.E. Kirkland. With the exception o f July 17th, when Kirkland collected Timmia austriaca on Spieden Island, all other collections were from Friday Harbor on San Juan Island. Even though the Marine Station could have provided the students access to a research vessel for inter-island travel, and commercial transportation was readily available, most o f the collection localities remained consistently the same from year to year. This was probably a result o f Frye's influence, and most o f the collecting was done either on San Juan Island or Orcas Island. Favorite spots on San Juan Island included Sportsman Lake, Trout Lake, Roche Harbor, Friday Harbor and on the new Station grounds on Point Caution. The Class o f 1925 had the first really good opportunity to thoroughly collect the Point Caution area as they  11  were the first students in a bryophyte class to be held at the new site. Common collection locations on Orcas Island included Eastsound, T w i n Lakes, Rosario, and one o f their favorite locations, M t . Constitution. Often their label information would be rather vague, such as the "north side o f Orcas Island", or "near Eastsound"; thus, it is difficult to relocate some o f their early collecting sites. In 1930 Frye collected throughout the year within the Friday Harbor area on Sah Juan Island. This was Frye's last year as the director o f the Station and a serious decline in the bryological activity within the islands, coincided with his departure. From this point on there was very little bryological collecting activity. O n July 4th, 1931, Ruth D . Svihla, a bryologist, visited and collected briefly on San Juan Island. In 1937 Elizabeth Young, Frye's daughter, collected at Friday Harbor. Presumably both o f these collections were the result o f short visits to the Station and did not represent formal research. Based on the herbarium data, Frye's last field work in the San Juan's occurred on July 1st 1944 when he collected on San Juan Island After 1944, most collecting represented casual trips to the area by students, although several well-known bryologists visited the area briefly. E l v a Lawton, according to her field notes, made a trip to the M t . Constitution area on July 22, 1956. While there, she collected Polytrichum piliferum,  Racomitrium  heterostichum, R. canescens var. ericoides, R. canescens  var. elongatum and Grimmia trichophylla.  Robert Ireland, a graduate student working under  the direction o f Lawton, visited Friday Harbor on M a y 19, 1962, and collected Grimmia trichophylla  var. muhlenbeckii.  Matia Island was visited by Larry Goodhew, Lawrence  O'Flaherty, Dale Durrwachter and James Gregory on February 18th 1962. Their visit may have represented part o f a class assignment as they collected extremely large quantities o f 18 species o f moss. It is possible that they may have been responsible for the extinction o f two species from the island. David Largent, another University o f Washington student, collected around the Station grounds on Oct. 23, 1965. W . B . Schofield visited San Juan Island briefly in 1970 and collected 12 mosses from around the ferry terminal area. 12  Mrs. Betty Higinbotham, a past President o f the American Bryological and Lichenological Society, had a summer home on San Juan Island. In a meeting with her in 1990 she indicated to me that she had never made extensive collections o f bryophytes within the islands. In 1985 she wrote a short article entitled "The Mini-Plant World o f Mosses and Lichens" which was published in the local San Juan Islands Almanac (Higinbotham 1985). In this article she lists some o f the most common species o f mosses and lichens but this list is not offered as a comprehensive survey o f the region's flora. According to the herbarium labels at Western Washington University, M r s . Higinbotham collected Racomitrium canescens var. ericoides from the M t . Dallas area, San Juan Island on A p r i l 4th, 1975. A recent search o f her bryological collections located at the Washington State University herbarium in Pullman revealed a fairly large number o f specimens still in the original collecting bags. Most o f the collections came from San Juan Island, but she also made trips to Lopez, Orcas, Shaw and Speiden Islands. Although she had indicated that she did very little collecting within the islands, her field work resulted in 61 species and 196 packets o f bryophytes. A l l o f these bryophytes were obtained during brief visits to their summer home between 1973 and 1984. Prior to the detailed exploration and collections represented by the thesis research reported here, essentially all bryological research done within the San Juan Islands has been the result o f casual or class collecting and there has never been a serious attempt to develop a comprehensive flora for the area.  Although the classes held at the University o f Washington  Marine Station provided many specimens, all came from frequently revisited locations and a systematic survey o f all o f the islands was never attempted.  Between 1858 and 1984  approximately 244 field days were spent in collecting bryophytes from only 10 of the islands found within San Juan County. The material gathered during this 126 year period has resulted in a species list o f approximately 137 species for the islands. O f these, a few species were collected that were not rediscovered during the present study. See Appendix A for a list of species not relocated during the present study. 13  These early collectors were fortunate to be able to do field work during the time when the islands were less disturbed by human activity than is the case today. Their collections hold historical significance in that many o f the areas that they repeatedly visited are now destroyed as sites for bryophytes or have undergone other major changes. One o f the best examples o f this is reflected in the early work done by G . B . R i g g on the peat resources and sphagnum bogs within the islands (Rigg and Richardson 1934). Although his work concerned palynological records and was not intended as floristic, he did map all o f the existing bogs; today most o f these bogs no longer exist (see Chapter 5 p. 75 for further details). In conclusion, although Frye and his students had a tremendous impact on bryological research within the San Juan Islands, they did not carry out a formal survey of the region. When Frye's work is combined with the earlier collection data, and the casual collections o f recent years, a reasonably large species list can be developed for the area. Yet all o f this earlier work was not sufficiently systematic to produce a bryophyte flora for the San Juan Islands.  14  CHAPTER 3 Description o f the Study Area  The San Juan Islands are part o f the San Juan archipelago which is located i n the northwestern part o f Washington State between Vancouver Island, Canada and mainland Washington (See Figure 1.). The center o f the area is located at 48 3 0 ' N latitude and 123 O'W longitude. These islands vary in size from less than one hectare (0.4 acre) to the largest at 14,744 hectares (36,432 acres). Elevation ranges from sea level to 734.4 m. (2,409 ft.) on the summit o f M t . Constitution. The archipelago includes 428 islands exposed at low tide, of these 175 are named. Together they represent a land mass o f 445.5 square kilometers (172 square miles) (Russell 1975). R o y M c L e l l a n , the first geologist to survey the San Juan Islands extensively, describes the islands as "In a restricted sense the San Juan Islands consist only o f the area included within San Juan County, Washington. T o the eastward occur islands belonging to Skagit and Whatcom Counties and these are commonly regarded as part o f the San Juan Island Group" (McLellan 1927). Although these islands are considered part o f the San Juan Island Group, for the purposes o f the present study only the Islands within San Juan County were surveyed. A discussion on the climate, geology, glacial history, soils, vegetation and Late Pleistocene and Holocene vegetation and climate for the islands found within San Juan County follows. Climate Good (1931) states "plant distribution is primarily controlled by the distribution of climatic conditions". Longton (1980) in a paper on the physiological ecology o f mosses comments; "temperature and availability o f water are shown to be significant i n controlling vegetative growth, while temperature and photoperiod appear to be involved i n regulation o f clearly defined seasonal cycles o f gametangial and sporophyte development". W i n d patterns also play an important role in the distribution o f moss spores (Zanten 1978).  15  Figure 1. Location map for the San Juan Islands.  Modified from Hebda (1983)  16  The climate o f a region, therefore, has a significant influence on the distribution patterns o f mosses The climate o f the San Juan Islands is characterized by relatively short, cool, dry summers and mild moderately wet winters, with 60% o f the rainfall occurring during the winter (Dietrich 1975). This climatic pattern results from the prevailing south southeast winter winds and the "rainshadow effect" created by the Olympic Mountains. During the summer months the semipermanent "Pacific High" that becomes established i n the area prevents storms from entering the region. Combined with the cool, southwest summer winds and the blocking o f the warm dry continental air mass by the Cascades, the summers are cool and dry. The San Juan Islands can be considered to be part o f the Mediterranean type climate found along the Pacific Coast of North America. Both Krajina (1959, 1965) and Chapman (1952) describe this region as within Koppen's Csb- Mediterranean type climate. This climatic region includes the southeastern portion o f Vancouver Island, across the Georgia Strait, north to Texada Island and along the mainland coast southward to the Lower Fraser Delta in British Columbia. Chapman (1952) cautions that there are significant local differences within this region; Krajina (1965) refers to this as "Csb-Mediterranean subhumid"). While there were only three official U . S . weather stations within the islands Diertrich (1975) used both official U . S . and nearby Canadian weather stations to prepare an isohyetal map o f estimated mean annual precipitation for the county, (Figure 2.). Weather data accumulated from the three official U . S . weather stations within the islands were also assembled by Phillips in his 1966 paper on the weather o f San Juan County, (Table 1.). Maunder (1968) describes seven patterns o f high pressure systems and nine patterns of low pressure systems that occur in the Pacific Northwest. The high pressure patterns usually dominate during the summer months, while the low pressure systems dominate the winter months. The wind patterns within the islands are predominantly from  17  Figure 2. Isohyetal map of the San Juan Islands.  Table 1. Temperatures and precipitation from known stations i n the San Juan Islands Location  Mean Daily Temperatures (F)  Precipitation (Inches)  Friday Harbor (San Juan Island)  Jan. == 7.4° C (45.4°) July == 23.9° C (75.1°)  Snow = 114.3 m m (4.5") Rain = 696.0 m m (27.4")  Olga (Orcas Island)  Jan. == 6.4° C (43.6°) July == 21.2° C (70.2°)  (7.7") Snow = 195.6 m m Rain = 731.5 m m (28.8")  Richardson (Lopez Island)  Jan. == not given July == not given  Snow = not given Rain = 500.4 m m (19.7")  based on Phillips (1966)  19  :  the south or southeast during the winter months, and from the west or northwest in the summers. Occasionally, during the winter seasons, l o w pressure systems form off the coast, thus large cold air masses from the Fraser River Canyon produce strong northeasterly winds. These winds are often destructive and result in large quantities o f blown down timber. During the summer months the winds are usually light and occur during the afternoon from the west. Three o f the sixteen patterns o f high and l o w pressure systems described by Maunder (1968) could have provided the right surface wind conditions necessary for the long distance dispersal of moss spores from the mainland. These three patterns are as follows: (1) a high pressure system is centered west o f central California/Oregon and a l o w pressure system is in the G u l f o f Alaska, (2) a l o w pressure system is centered between 45 and 50 N latitude and finally, (3) a low pressure area is centered over the G u l f o f Alaska. A l l three o f these patterns result in a southwesterly to westerly air flow that could have carried moss spores into the islands. See the discussion under phytogeography for additional details i n Chapter 6, p. 81.  20  Geology and Soils The composition and physical nature o f the substratum plays an important role in the distribution o f mosses (Richardson 1981). Some species such as Racomitrium lanuginosum, R. heterostichum, R. elongatum, Andreaea rupestris and A.  megistospora  are restricted to acidic substrata, while others such as Eucladium verticillatum,  Crumia  latifolia and Conardia compacta are confined to alkaline or calcareous substrata. The geology and soils o f a region significantly influence the distribution o f the mosses. M c L e l l a n (1927) comments that "the San Juan's probably represent the down-turn of an ancient piece o f orogenic crusts most o f which was long since consumed". In a more recent work Danner (1966) proposes that "the San Juan Islands compose the older core o f an ancient mountain system that once extended across the Pacific Northwest from Vancouver Island into Central Washington". Cowan (pers. comm. 1994) hypothesizes that these islands are composed o f terranes that formed around 90-100 million years ago that subsequently they collided with the Wrangellian terrane. According to the paleomagnetic data it appears that these terranes may have formed near the latitudinal position o f Baja California (Brandon et al. 1988). While there is debate as to where the terranes collided, Cowan (pers. comm. 1994), feels that the collision took place off the Baja California coast and that the resulting terranes migrated northward until they reached their current position about 55 million years ago. Thus, by the early Tertiary Period, these islands were in their current position. Brandon et al. (1983, 1988) describe fourteen lithologic units within the islands. These rock units range from Early Paleozoic to Quaternary i n age and can be divided into four groups. First, the San Juan Thrust System is composed o f a series o f thrust faults and nappes ranging in age from the early Paleozoic to the Late Cretaceous. It includes the following nine units: Constitution Formation, Deadman Bay Volcanics, East Sound Group, Fidalgo Complex, Garrison Terrane, Lopez Structural Complex, L u m m i Formation, Orcas Chert, and the Turtleback Complex. The rocks within these units were  21  subjected to Late Cretaceous thrusting and high-pressure metamorphism. Thus they represent some o f the oldest bedrock material in the Islands. The second group, the External Units, were formed during the Late Triassic to Early Cretaceous and may or may not have been subjected to the Late Cretaceous thrusting. They also did not undergo the high-pressure metamorphic conditions that affected the earlier units. The following three units are included i n this group: Haro Formation, Nanaimo Group, and the Spiedon Group. These units occur either in front o f or below the San Juan Thrust System. Third, the Post Orogenic Unit group contains only the Chuckanut Formation, an Eocene deposit composed o f nonmarine sandstone and conglomerate. This formation is exposed only on Patos, Sucia, Matia and Puffin Islands, where it replaces the Nanaimo Group. The fourth group contains only the Quaternary Cover, which is composed o f glacial till and sediments that were deposited during the Pleistocene and Recent Epochs. While Brandon (1988) does not break this group into individual units, M c L e l l a n (1927) divides it into four formations: the C o l wood Formation, composed o f sediments and alluvium deposited during the Recent Epoch; the Vashon and Admiralty Formations, composed o f glacial tills and sediments that were deposited during the Pleistocene; and finally the Puyallup Formation, composed o f interglacial sediments deposited during the Pleistocene. These deposits range i n size from clay particles to gravely till and can be up to 45.7 m (150 ft.) thick in some areas. The Quaternary Cover deposits are also responsible for the abundant sand spits and tombolas found throughout the islands. Thus, these older structures have provided a wide variety o f rock types that can be found among the islands. Figure 3. shows the locations, and Table 2. contains further descriptions o f these formations. While a large variety o f rock types occur within the islands, the limestone deposits found in the Deadman Bay Volcanics and East Sound Group (on Orcas and San Juan Islands) are the most significant geological features with respect to the bryophytes. When these limestone deposits occur associated with fresh water, an unusual habitat is es 22  Figure 3. Geology of the San Juan Islands. raw  ^aw  R Z W  R 1 W  738*/  K  7377V  T38N 737«  s  \  T3SN T3SN  I  I  _73*V  T34N  O  QUATERNARY COVER CHUCKANUT FORMATION NANAIMO GROUP HAROTERRANE  (_) 0ft  . . (£3 C O N S T I T U T I O N  FORMATION  SPIEDEN GROUP HARO FORMATION  0  LOPEZ STRUCTURAL COMPLEX  DECATUR TERRANE  (T)  GARRISON TERRANE  LUMMI FORMATION FIDALGO COMPLEX  DEADMAN BAY TERRANE f~)  ORCASCHERT DEADMAN BAY VOLCANICS  TURTLEBACK TERRANE EAST SOUND GROUP TURTLEBACK COMPLEX  Based on Brandon et al. (1988)  23  o f0  J 5 kilometers  5 miles  W30-  -i  Table 2. Geologic Formations i n the San Juan Islands Tectonic Group RECENT  Name o f Unit QUATERNARY COVER  Age Pleistocene & Recent  POST OROGENIC  CHUCKANUT FORMATION  Eocene  THE EXTERNAL UNITS  NANAIMO GROUP  Late Cretaceous  HARO TERRANE Haro Formation Spieden Group THE SAN JUAN THRUST SYSTEM  Lithologic Description glacial till and sediments Nonmarine sandstone and conglomerate Marine and nonmarine sandstone, conglomerate, and shale  Late Triassic  marine volcaniclastic sandstone and conglomerate with minor silicic tuff Late Jurassic & marine sandstone and Early Cretaceous conglomerate derived from an arc-volcanic source  DECATUR TERRANE Lummi Formation Fidalgo Igneous Complex LOPEZ STRUCTURAL COMPLEX CONSTITUTION FORMATION GARRISON TERRANE  DEADMAN BAY VOLCANICS Orcas Chert Deadman Bay Volcanics TURTLEBACK TERRANE East Sound Group Turtleback Complex  Latest Jurassic clastic marine sequence & Early Cretaceous Middle & Late An ophiolite with younger Jurassic superimposed volcanic arc Late a fault zone, with sandstone, Cretaceous pebbly mudstone, pillow lava and chert Jurassic or volcaniclastic sandstone, with Early Cretaceous imbedded mudstone, ribbon chert, pillow lava, and green tuff Permian to high-pressure metamorphic Early Triassic rocks, of metamorphism mafic schist with minor quartzmica schist, greenschist to albite-epidote amphibolite facies Triassic & Early Jurrassic Early Permian to Triassic  ribbon chert, minor pillow basalt and limestone pillow basalt with minor interbedded Tethyan-fusulinid limestone and ribbon chert  Eearly Devonian an arc-volcanic sequence with to Early Permian minor interbedded limestone, with non-Tethyan fusulinids probably a plutonic complex consisting of Cambrian tonalite and subordinate gabbro  Based on Brandon et al. (1988)  24  established that supports a variety o f calcicolous mosses. Tufas, formed by the precipitation o f C a C 0 3 on the surfaces o f calcicole mosses can often be found in such habitats (Flugel 1982). The limestone deposits are also o f interest because they do not occur extensively in the adjacent islands or on the mainland. Thus the tufa formations that occur within the islands represent distinctive habitats. These limestone deposits have been quarried extensively, and while they contributed significantly to the island's early economy, they have been badly disturbed as a consequence. Therefore the distinctive calcareous habitats are now somewhat limited in the islands. During the Pleistocene Epoch the islands were extensively glaciated. Easterbrook (1969) has documented three periods o f glaciation and associated interglacial periods. The Stuck Glaciation period followed by the short Puyallup Interglacial began around 70,000 years B . P . Approximately 40,000 years B . P . the Salmon Springs Glacial period began, persisting until about 28,000 years B . P . This was followed by the Olympia Interglacial which persisted until about 19,000 years B . P . During these early periods o f glaciation the islands were completely covered by thick deposits o f ice or were submerged by marine water. The final episode o f glaciation began around 19,000 years B . P . and has been called the Fraser Glacial period. During this period o f glaciation the Puget and Juan de Fuca Lobes advanced southward at different rates o f speed. The Juan de Fuca Lobe advanced across the San Juan and adjacent islands as well as the southern tip o f Vancouver Island, ultimately ending in the western Strait o f Juan de Fuca about 17,000 years B . P . While the Juan de Fuca Lobe was advancing, the Puget Lobe had already pushed south o f Seattle by around 15,000 years B . P . (Waitt & Thorson 1983). Easterbrook (1969) divided the Fraser Glacial period into the following:  The Vashon  Stade, a period o f glacial advance that occurred between 19,000 - 13,000 years B . P . and covered the top o f M t . Constitution with a layer o f ice that was approximately 915 - 1097 meters (3000 - 3600 ft.) deep; The Everson Interstade, a time o f glacial retreat between 25  13,000 - 11,000 years B.P.; and finally, The Sumas Stade, (11,000 - 10,000 years B.P.) a small advance o f ice that probably did not go very far beyond the United States - Canada border, see Table 3. Recent research by Dethier et al. (1995) states that "As the Fraser ice retreated, marine waters invaded the northern Puget Sound and adjacent areas o f British Columbia, inundating most o f the San Juan Islands  Abundant invertebrate fossils representing  cold water species with cool-temperate to Arctic affinities indicate that these marine waters were cold. The macrofossils further suggest that this marine environment was similar in temperature to that which is more characteristic o f the northern G u l f o f Alaska at the present time. Dethier et al (1995) also suggest that the Sumas Stade had no significant influence on the climate. Thus, by about 13,500 years B.P. the higher regions of the islands (such as M t . Constitution, Orcas Knob, and Dallas M t . ) were probably above water, ice free, and were surrounded by a cool marine environment (see Figure 4.). A s the glaciers retreated, large quantities o f outwash and interglacial sediments, composed o f various types o f sand and gravel, were left on the islands. Glacial till, a hard compacted sediment, was deposited over both the earlier sediments and on bedrock. This material, known as Vashon T i l l , covers most o f Lopez and Decatur Islands and is composed o f all sizes o f lithic material, including clay. O n the southwest side o f Lopez Island the depth o f the till was measured at 16.1 m (53 ft.) (Russell 1975). Both M c L e l l a n (1927) and Russell (1975) note that large granite boulders, (called glacial erratics) were deposited along with this till and can be found scattered throughout the islands. The final glaciomarine drift deposits were left on top o f the earlier deposits as the Fraser Ice retreated. This fossil-rich material is widespread and probably is 10-15 feet thick (Russell 1975). Schlots et al. (1962) describe four types o f soil associations found throughout the islands (see Figure 5., and Tables 4. and 5.). The Bellingham-Coveland-Bow association, composed o f "poorly drained to imperfectly drained soils o f the basins and l o w glacial till 26  Table 3. Glacial history i n the Puget Trough. Yr. B.P. 0  Geological / Climatic Units  1000 Cool and Moist  Event  Quercus pollen decreases and Thuja-Chamaecyparis pollen increases. (L. Heusser 1983)  2000 3000 4000 5000 6000 7000 8000  H Y P S I T H E R M A L  Garry Oak-Douglas Fir Subzone is well established in the Saanich Inlet, on Vancouver Island. (L.Heusser 1983) Warm and Dry Mt. Mazama Eruption  9000 Cool and Moist 10000 SUMAS STADE 11000 12000  EVERSON INTERSTADE Cool and Dry  13000  Vashon Ice retreats rapidly north of islands, stops at US/CAN border. (Waitt et al. 1983) In the San Juan Islands, higher areas were above water and ice free. (Deither et al. 1995)  14000 15000 VASHON STADE  Puget Lobe advances to the Seattle area, then begins to retreat. (Waitt et al. 1983)  16000 17000 Alpine Glaciers greatly diminished. (Waitt et al. 1983) 18000  19000  Juan de Fuca Lobe advances to the western strait of Juan de Fuca. (Waitt et al. 1983)  20000  Alpine Glaciers at maximum positions. (Waitt et al. 1983)  21000 22000  OLYMPIC INTERGLACIAL  23000 (Easterbrook 1969)  27  Figure 4. Marine Limit at about 13,500 yr. B.P., submerged area is shaded.  28  Figure 5. Soils in the San Juan Islands.  Based on Scholts et al. (1957)  29  Table 4.  Soil types found in the San Juan Islands (after Schots et al. 1962)  1. Bellingham - Coveland - Association a. Bellingham Soils  fairly large areas  b. Coveland Soils  less extensive in the area  c. B o w Soils  most extensive in the area  d. Norma Soils  few small areas  e. Semiahmoo Soils  few large areas, and a number of smaller areas  f. Tanwax Soil g. Orcas Soils h. Roche-Rock Outcrop Complex  ' small but prominent areas  2. Roche - San Juan Association a. Roche Soils fairly extensive areas b. San Juan Soils c. Rock land large areas d. Alderwood Soils  fairly large areas  e. Everett Soils  small isolated areas  f. g. h. i.  Indianola Soils Pickett Soils Coveland Soils Semiahmoo Soils  3. San Juan - Everett Association a. San Juan Soils small areas b. Everett Soils small areas c. Indianola Soils small areas d. Roche Soils e. Alderwood Soils 4. Pickett - Rock land Association a. Pickett Soils are dominant b. Rock land c. Semiahmoo Soils  small isolated areas  d. Orcas Soils e. Tanwax Soils  30  Table 5. Soil units in the San Juan Islands SOIL UNIT Alderwood  SUBSOIL  SURFACE T E X T U R E gravely or stony, medium texture  cemented till  Bellingham  clay or silt loam  clay or silt loam  Bow  silt loam  clayey  Coveland  silt loam  Everett  clayey, clay loam, sandy jdayjoam loose, gravelly, sandy, gravely or stoney moderatly coarse texture porous  Indianola  moderatly coarse stoney, little gravel  loose, gravelly, sandy, porous  Norma  loam  sandy loam, clay loam or clay  Orcas  moss peat, bog  pale-brown, spongy fibrous peat  Pickett  medium textured and stoney  sandstone bedrock  Roche  brown, medium textured may be gravely or stoney stones, 12-24 inches, on surface rock outcrops of basalt, sandstone & argillite dark-colored, moderatly coarse textured, gravelly, or stony muck, derived from sedge peat finely divided sedimentary peat  dense, slowly permable glacial till  Roche-Rock Rock Land San Juan  Semiahmoo Tanwax  ? bedrock coarse textrued & cemented moderatly fine textured glacial till sedimentary peat finely divided organic material including alage, excreta from marine animals  based on Schlots et al. (1962)  31  NATIVE VEGETATION doug fir, lodgepole pine, willow, alder, big leaf maple, willow red cedar, hemlock, alder, big leaf maple doug fir, hemlock, alder, big leaf maple, salal, Oregon grape, ocean spray probably was grass and sedges doug fir, lodgepole pine, madrona, salal, Oregon grape, sword fern, moss doug fir, lodgepole pine, red cedar, alder, big leaf maple, salal, oregon grape, ocean spray, sword fern doug fir, sitka spruce, white fir, lodgepole pine, willow, ocean spray, bracken fern, sedges sphagnum, hypnum moss, spirea, lodgepole pine doug fir, red cedar, lodgepole pine, a few madrona doug fir, lodgepole pine, willow, alder, salal, oregon grape grass little cover, soils are too shallow grasslands  sedges reed canary grass  plains on glaciated uplands". The topography o f this association is nearly level to gently sloping, and it occurs on most o f the larger islands. The Roche-San Juan association is composed o f "dominantly moderately well drained to well drained soils o f the glacial till plains and rocky uplands". This is the largest association in the area and it is characterized by a predominantly gently rolling to rolling topography that covers most o f San Juan Island. The San Juan-Everett association is characterized by "somewhat excessively drained soils on glaciated uplands and outwash plains". Very small examples o f this association are found throughout the islands, mainly in rolling to steep areas on Waldron and San Juan Islands. Finally, the Pickett-Rockland association is composed o f "dominantly well drained soils on uplands". It occurs on fairly steep, irregular, rolling to steep or precipitous topography, and is mostly found on Orcas and Blakely Islands.  32  Vegetation  The vascular plant vegetation influences the distribution o f mosses by providing substrata and microhabitats. This is especially evident i n the epiphytic mosses, which are often restricted to certain tree species. Therefore, an overview o f the vascular plant flora is given below. Franklin and Dyrness (1973) place the San Juan Islands i n the Puget Sound Area of the Tsuga heterophylla Zone. The Puget Sound Area is unusual because it is composed o f plant communities that differ from most o f the Tsuga heterophylla Zone, and they often do not occur anywhere else in western Washington. Several of these plant communities are found within the San Juan Islands, and include populations of Juniperus scopulorum (Rocky Mountain juniper) on open rocky slopes near salt water, stands o f Populus tremuloides (quaking aspen) in riparian areas on glacial till soils, and clusters o f Betulapapyrifera  (paper birch) i n stream valleys and  riparian areas on San Juan and Orcas Islands. Using Krajina's (1959) classification system the San Juan Islands are in the Coastal Douglas Fir Zone. This temperate Mediterranean coniferous forest is a response to the rain shadow effect generated by the Olympic Mountains. Although this zone is dominated by Pseudotsuga menziesii, the following species are also present: Abies grandis (grand fir), Arbutus menziesii (madrona), Pinus contorta (lodgepole pine), Thuja plicata (western red cedar), Tsuga heterophylla (western hemlock), Gaultheria shallon (salal), Holodiscus discolor (ocean spray), Berberis nervosa (Oregon-grape), and Polystichum munitum (sword fern). Krajina (1965) divides this zone into two subzones: The drier Garry Oak- Douglas F i r subzone, composed o f the following species; Quercus garryana (Garry oak), Camassia quamash (camas), Collinsia grandijlora (large-flowered collinsia), Plectritis congesta (sea blush), and Zygadenus venenosus (poison-camas); The wetter Madrono-Douglas F i r subzone, composed o f the following species; Arbutus menziesii (madrona), Pinus contorta (lodgepole pine), P. monticola (western white pine), Thuja plicata (western red cedar), Gaultheria shallon (salal),  33  Holodiscus discolor (ocean spray), Berberis nervosa (Oregon-grape), and Polystichum  munitum  (sword fern). Atkinson and Sharpe (1985) make several interesting comments concerning the vascular flora o f the San Juan Islands: (1) "the absence or scarcity o f many species typical o f wet coniferous forest zone is a striking element o f the San Juan Island's". These missing or scarce species include; Acer circinatum (vine maple), Rhododendron macrophyllum  (Pacific  rhododendron), Achylys triphylla (vanilla-leaf), Rhamnus purshiana (cascara), and Vaccinium ovatum (evergreen huckleberry). (2) The flora is "host to a number o f taxa usually associated with dry habitats east o f the Cascades". Some o f the 21 eastern taxa are Juniperus (Rocky Mountain juniper), Woodsia scopulina (Rocky Mountain woodsia), Opuntia  scopulorum fragilis  (prickly pear cactus), Poa scabrella (pine blue grass), and Camissonia contorta (contorted-pod evening-primrose). They relate the first two observations to the dry conditions brought about by the rain shadow effect o f the Olympic mountains. Finally (3). "the presence o f a relic late glacial flora on the summit o f M t . Constitution, where several subalpine and northern species linger". A m o n g these species are Athyrium distentifolium (alpine lady-fern), Aster sibiricus (arctic aster), Epilobium alpinum (alpine willowherb), Erigeron trifidus (dwarf mountain daisy), and Saxifraga bronchialis (matted saxifrage). While the San Juan Islands are classified as being i n the Coastal Douglas Fir Zone, and have a good representation o f plants from both o f the subzones, the M t . Constitution area still retains some o f the unusual and distinct subalpine - alpine elements, possibly surviving from the time when the site may have been first colonized. Many o f the moss species found during the present study exhibit these similar distribution patterns. The discussion under phytogeography provides further details.  34  Late Pleistocene and Holocene Vegetation and Climate  The importance o f the relationship between temperature and moisture to the distribution of vegetation has been clearly demonstrated by Krajina, (1959, 1965) and Franklin and Dyrness, (1973). The fossil arboreal and non-arboreal pollen found in peat deposits can be used to reconstruct and infer past changes in climatic and vegetation patterns (Pielou 1991). During the Late Pleistocene, from approximately 14,000 yr. B . P . to 10,000 yr B.P., coastal Washington and Southern British Columbia were covered predominately with Pinus contorta and Alnus sp., and the climate was cool and moist (C. Heusser 1960, L . Heusser 1983, Hebda 1983, Barnosky 1984, 1985, Mathewes 1985). It was during this cool moist period around 13,500 yr. B . P . that the higher areas in the San Juan Islands were ice free and also exposed above the cool marine waters (Deither et al. 1995). Thus plants could have re-colonized the Islands as early as 13,500 yr. B . P . Beginning around 10,000 yr. B . P . a warm, dry period began in the Puget Trough, (Hibbert 1979, Barnosky, 1981, 1984, Mathewes & Heusser 1981, L . Heusser, 1983, Mathewes 1985). This hypsithermal period lasted until around 3000 yr. B.P., and was characterized by the appearance o f Quercus, and Pseudotsuga pollen and Pteridium spores i n the palynological record. A t the same time there was a decrease in the amount o f pollen from Pinus and other temperate species. This hypsithermal event is well documented i n the marine core samples from the Saanich Inlet on Vancouver Island. L . Heusser (1983) found that large quantities o f Quercus pollen had been deposited during this period and comments that Krajina's (1965) Garry OakDouglas F i r Subzone was well established in the Sannich Inlet by about 7000 yr. B.P.. Following this warm period, about 3000 yr. B.P., an increase in Tsuga heterophylla and Thuja type pollen indicates that a moist cooling trend had begun in the Puget Trough, (Hibbert 1979, Barnosky 1981, 1985, Hebda 1983, L . Heusser 1983).This cool moist climate has persisted throughout the region today (see Table 6).  35  Little palynological research has been done within the San Juan Islands. Hanson (1943) examined the pollen deposits in two Orcas Island bogs, one near the summit o f M t . Constitution and the other a lowland bog at Killebrew Lake.  He states that i n both bogs, "lodgepole pine was  the predominant species when the lowest pollen-bearing sediments were deposited". The large numbers o f Pinus contorta (lodgepole pine) now found i n the Orcas Island bogs are not unusual because Pinus contorta was an early dominant pioneer species within the Puget Trough. Although Hanson (1943), did not date his pollen cores, he did note a layer o f ash in all o f the bogs that he studied. In the M t . Constitution bog, the ash layer was found between 7.0 and 7.5 meters, and in the Killebrew Lake bog the layer occurred around 7.0 meters. Hanson (1943) attributes this ash layer to an eruption o f Glacier Peak, but more recent work by Powers (1964) and Sarna-Wojcicki et al. (1983) found this ash layer to have originated from M t . Mazama. This provides the ash layers in the bogs a date o f about 6700 yr. B . P . A t about 6700 yr. B . P . the pollen deposits from the M t . Constitution bog indicate that the surrounding vegetation was composed mostly o f Pinus contorta and Pinus monticola, with moderate amounts o f Abies grandis and Picea sitchensis.  Both Pseudotsuga menziesii and Tsuga heterophylla were poorly  represented at this time. In the Killebrew Lake bog the dominant tree species were, Pseudotsuga menziesii and Tsuga heterophylla, with minor amounts o f Pinus contorta, Abies grandis and a trace amount of Picea sitchensis pollen. During his study Hanson (1943) omitted the deciduous trees, herbs, and grasses because they were poorly represented i n the pollen deposits. It is difficult to determine accurately when Quercus entered the islands. Using L . Heusser's (1983) reports based on nearby Sannich Inlet, i f the Garry Oak-Douglas F i r Subzone was well established by 7000 yr. B.P., it is reasonable to assume that it would also be well established in the San Juan Islands at approximately the same time. Hanson's (1943) Killebrew Lake bog pollen records show a marked increase in Pseudotsuga menzesii pollen levels coinciding with a decrease i n the Pinus contorta and Pinus monticola pollen around the time o f the M t . Mazama ash layer (6700 yr. B.P.). Correspondingly, the appearance o f Pseudotsuga menziesii pollen in the M t . Constitution bog does not occur until 36  Table 6. Pollen records in the Puget Trough yrB.P  Bear Cove. GeologicalClimatic Unit Vane. Isl.  Sannich Inlet, Vane. Isl.  0.0  Thua type Tsuga heterophyllaCupressaceae (Thuja)  1000  moister & cooler  San Juan Isis., W A Kille. Lake Pseudotsuga Tsuga heterophylla  Mt. Const. Pinus contorta Pseudotsuga menzesis Tsuga heterophylla  Nisqually Lake, W A  Pseudotsuga -TsugaThuja Forest  Battle Gr. Davis Lake, W A Lake, W A  Pseudotsuga -TsugaThuja forest  cool & moist  Pseudotsuga -TusgaThuja Forest Cooler & Moist  cool & moist  2000  Temperate & wet  3000 Garry OakDoug Fir Subzone  HYPSITHERMAL  Tsuga heterophyllaPicea  4000  well established Pseudotsuga - A In usThuja Forest  warmer & drier  Moderate warm & dry 5000  6000 MT. MAZAMA ASH LAYER PiceaPseudotsguaAlnusPteridium  7000  AlnusPteridium  Pinus contorta  Pseudotsuga menziesi  P. monticola  Tsuga hetrophylla  Maximum warmth & dryness  warmer & wetter  Oak Savannah & Pseudotsuga -Alnus Woodland Warm & Dry  warmer & drier  8000  Pseudotsuga -Alnus Woodland  Oak Savannah Pseudotsuga -Pine Woodland  9000 Picea-Tsuga mertensianaAlnus cool & moist  10000  Pinus cool& moist  SUMAS STADE  11000  Pinus Parkland  Cool & Wet mixed Pinus - Abies Woodland  Picea-PinusTsuga Woodland  Pinus-Alnus EVERSON 1NTERSTADE  cool & dry  cool & moist Picea-Tsuga Woodland  12000  Picea-Tsuga Woodland 13000 VASHON STADE  High Points Ice Free &  Pinus Woodland  Above Water  Cool & Dry  Glacial Ice  14000  Plcea-Pinus Parkland Tundra 15000  UNNAMED INT ERST ADE  Picea Parkland Tundra  Plcea-Pinus Parkland  16000 Easterbrook (1969)  Hebda (1983)  Heusser (1983)  Hanson  Hanson  Hibbert  Barnosky  (1943)  (1943)  (1979)  (1985)  Waittetal (1983) 37  Barnosky (1985)  just above the ash layer; while the Pinus contorta level remains fairly high, it does show a small decrease. The Pinus contorta levels have probably remained high throughout the M t . Constitution bog profile as a result o f the unusual edaphic condition in the area (Hanson 1943), and therefore may not have responded to the overall changes i n climate. Since pine tends to be a prolific pollen producer, and would have been near the area o f deposition, it is likely to be overrepresented in the profile and reflect local conditions rather than the general vegetation o f the island. The cooling, moist trend that began around 3000 yr. B . P . is well documented in the pollen records by the increase o f Thuja type and Tsuga heterophylla pollen, (L. Heusser 1983, Hebda 1983). Although Thuja type pollen was not recorded by Hanson (1943) i n the Orcas Island bogs, he does show the distribution o f Tsuga heterophylla.  In the Killebrew Lake pollen profile, Tsuga  heterophylla pollen appears throughout the entire depth o f the bog. It reaches it's highest concentrations just below the Mazama ash layer (6700 yr. B.P.) and remains relatively high throughout the rest o f the profile. The M t . Constitution bog shows a different pattern, here the Tsuga heterophylla pollen starts to appear in small amounts above the ash layer and reaches its highest concentrations at the two meter depth. In both bogs Tsuga heterophylla does not become the dominant species. It would appear that the dry Mediterranean type climate has prevented both Tsuga heterophylla and Thuja plicata from becoming well established within the islands. The vegetation patterns in the islands today reflect the dry Mediterranean type climate that occurs within the rain shadow o f the Olympic Mountains. So little palynological work has been done in the islands that caution must be used in trying to interpret the patterns o f vegetation succession. Additional palynological research would result in a better understanding o f the earlier patterns o f vegetation and might show when and where Quercus first arrived in the islands. Since there are pollen profiles for most o f the Puget Trough, it would be interesting to see just how the San Juan Islands fit into the overall picture.  38  CHAPTER 4 Methods and Materials  The flora discussed here is based on collections o f mosses made over a period o f four years, from 1990 to 1993. A small number o f mosses were collected during a brief visit to the islands in 1988. The Islands were visited during every month o f the year except December and January. Historical collections made by earlier botanists were examined from the following herbaria: University o f British Columbia, ( U B C ) , University o f Washington, ( W T U ) , Washington State University at Pullman, (WS), Western Washington University, ( W W B ) , Portland State University, ( H P S U ) and the herbarium at the Friday Harbor Laboratories, ( F H L ) . These appear to represent most o f the historically significant collections. Permission to collect specimens on private property was obtained from numerous owners as well as the following agencies: the U . S . Fish and Wildlife Service, Refuge Islands; the U . S . National Park Service, San Juan Island National Historical Park; The Nature Conservancy; The San Juan Preservation Trust; Washington State Parks, the University o f Washington, and Seattle Pacific University. Transportation to and from some o f the islands was provided by the U . S . Fish and Wildlife Service, The Nature Conservancy, the U . S . M a i l Boat, and Washington State Parks Department. The larger islands were accessed from the Washington State Ferry system. A total o f 28 islands were visited, resulting in 6021 collections made from 159 sites. See Table 7. and Figure 6. provide details o f the collecting locations. Prior to visiting an island a careful literature search was done to determine the locations o f any unusual features, i.e. caves, limestone outcrops, etc. Species that are often associated with these types o f sites, as well as the common ones, were collected. Historical locations were also visited whenever the label information was sufficient to locate the old collecting site.  39  Table 7. Collecting locations. No. 1 2 3 4  Island Patos Patos Patos Sucia  5 6  Sucia Sucia  7  Sucia  8 9 10 11 12 13 14 15 16 17  Matia Matia Clark Orcas Orcas Orcas Orcas Orcas Orcas Orcas  18  Orcas  19  Orcas  20  Orcas  21  Orcas  22  Orcas  23  Orcas  24 25  Orcas Orcas  26  Orcas  27 28 29 30  Orcas Orcas Orcas Orcas  31  Orcas  32  Orcas  33  Orcas  34 35  Orcas Orcas  Location West end of island Middle of Island East end of island N.side of Echo Bay & Ewing Cove area Shallow Bay & end of Echo Bay Service roads & mixed woodland area Mud Bay, Fossil Bay, Johnstone Pt, & E V . Henry Pt. West end of Island Middle & East end of Island entire island Eagle Lake area Along Doe Bay Road Doe Bay Resort Obstruction Pass area and C.G. Olga Doe Bay Cemetary Entrance Mountain area East Sound Beach & Palisades area Rosario Resort & trail to Cascade Lake E. end of Cascade Lake, & Sunrise Rock area Cascade Lake loop trail, west of the foot bridge Trail to Cascade Lake from Cold Springs Along the road next to Cascade Lake Envin. Learning Center, & trails to Cascade Falls Doe Bay Rd. by the water tanks Trail to Cascade Falls from Mountain Lake Mt. Lake Loop Trail, below the dam, & dirt road junction Mt. Lake Loop Trail Mt. Lake Campground area Road up to Mt. Constitution Trail from Mt. Constitution to Cold Springs Trail from Cold Springs to Mt. Lake Trail from Cold Springs to Mt. Lake Mt. Constitution and Summit Lake area Bonnie Sliger Memorial Trail Mt. Lake Loop Trail  T38N T38N T38N T38N  T. R. S. R2W Sec. R2W Sec. R2W Sec. R2W Sec.  17 16 15 23,24  Lat. & Long. 48 47'N 122 57'W same as above same as above 48 4 5 ' N 122 52'W  T38N R2W Sec. 23 T38N R2W Sec. 23  same as above same as above  T38N 26 T38N T38N T37N T37N T37N T36N T36N T36N T36N T36N  R2W Sec. 25 &  same as above  R l W S e c . 29 R l W S e c . 28 R l W S e c . 13 R l W S e c . 25,26 R l W S e c . 35 RlWSec. 2 R l W S e c . 16 RlWSec. 9 RlWSec. 8 RlWSec. 6  48 4 5 ' N 122 50'W same as above 48 4 2 ' N 122 45'W 48 36'N 122 46'W 48 38'N 122 47'W 48 38'N 122 47'W 48 37'N 122 50'W 48 3 8 ' N 122 50'W 48 37'N 122 50'W 48 38'N 122 51'W  T37N RlWSec. 31  48 38'N  122 52'W  T37N R1W Sec. 32  48 38'N  122 51'W  T37N R l W S e c . 31 T37N R l W S e c . 29 & 32 T37N R l W S e c . 31 T37N R l W S e c . 32 & 33 T36N R l W S e c . 4 T37N R l W S e c . 33  same as above 48 4 0 ' N  122 50'W  48 39'N  122 50'W  same as above 48 38'N 48 39'N  122 50'W 122 50'W  T37N R l W S e c . 33 & 34 T37N R1W Sec. 34 T37N R l W S e c . 33 T37N R l W S e c . 33 T37N R l W S e c . 21, 28, 29 T37N R l W S e c . 29  48 40'N  122 49'W  T37N R l W S e c . 20  same as above  T37N R l W S e c . 21  48 40'N 122 50'W  T37N R l W S e c . 21, T37N R1W Sec. 27  48 37'N 48 40'N  40  48 40'N 122 48'W same as above 48 39'N 122 50'W 48 4 0 ' N 122 50'W 48 40'N  122 50'W  122 50'W 122 48'W  Location Mt. Picket area, & South Ridge Trail Buck Mountain area Buckhorn Road to Racoon Point Day Lake area Buckhorn Area Road into the Day Lake & Buck Mt. area East Sound, below Oyster Ship Resturant Madrona Point Park East Sound Point Doughty Beach between Pt. Doughty and Camp Orkila entire island Camp Orkila Enchanted Forest Road Woodlawn Cemetary Old Red Q Rock Quarry Turtleback Road Turtleback Road West Crow Valley Rd. near jet. of Nordstrum Lane Turtleback Road Orcas Knob  No. 36  Island Orcas  37 38 39 40 41 42  Orcas Orcas Orcas Orcas Orcas ck Orcas  43 44 45 46  Orcas Orcas Orcas Orcas  47 48 49 50 51 52 53 54  Freeman  55 56  Orcas Orcas  57  Orcas  58  Orcas  59 60 61 62 63 64 65  Orcas Orcas Orcas Orcas Orcas Orcas Orcas  66 67 68 69 70 71 72 73 74 75 76  Orcas Orcas Orcas Orcas Orcas Orcas Jones Waldron Waldron Waldron Waldron  Along Deer Harbor Rd. across from Massacre Bay Mt. Woolard Area, along main road Mt. Woolard Area Martin Lake Mt. Woolard Area, Ladd Lake Killebrew Lake Guthrie Cove Road Road to Grindstone Bay Montfort Prop. & Roads from Mt. Woolard area to Grindstone Bay Area around the Ferry Terminal Pole Pass Road Area Camp Four Winds Spring Pass Develp. Loop Road Spring Pass Road Public Beach No. 240 Entire Island Point Disney Between Pt. Disney & Mail Bay Mail Bay Point Hammond Area  77 78  Waldron Waldron  Along the road by the School Road to the Cemetary  Orcas Orcas Orcas Orcas Orcas Orcas Orcas  T.R.S. T37N R l W S e c . 27, 34  Lat. & Long. 48 40'N 122 47'W  T37N T37N T37N T37N T37N  17 17 17 7 18  48 41'N 48 4 2 ' N 48 41'N 48 4 2 ' N 48 44'N  122 51'W 122 51'W 122 50'W 122 52'W 122 52'W  T37N R2W Sec. 25  48 42'N  122 53'W  T37N T37N T37N T37N  48 41'N 48 43'N 48 4 3 ' N 48 43'N  122 54'W 122 55'W 122 57'W 122 57'W  RlWSec. R1W Sec. RlWSec. RlWSec. RlWSec.  R2W Sec. 14 R2W Sec. 11 R2W Sec. 9 R2W Sec. 9  T37N R2W Sec. 16 T 3 7 N R 2 W Sec. 15,16 T37N R2W Sec. 15 T37N R2W Sec. 21 T37N R2W Sec. 20 T37N R2W Sec. 29 T27N R2W Sec. 28 T37N R2W Sec. 33  48 4 2 ' N 122 57'W 48 41'N 122 57'W 48 4 2 ' N 122 56'W 48 40'N 122 57'W 48 4 1 ' N 122 57W 48 4 0 ' N 122 57'W same as above 48 39'N 122 57'W  T37N R2W Sec. 32 T37N R2W Sec. 29, 30,31,32 T36N R2W Sec. 5  48 40'N 48 4 0 ' N  122 57'W 122 58'W  48 37'N  122 58'W  T36N R2W Sec. 10  48 37'N  122 55'W  T36N R2W Sec. T36N R2W Sec. T36N R2W Sec. T 3 6 N R 2 W Sec. T 3 6 N R 2 W Sec. T36N R2W Sec. T36N R2W Sec.  11 12 14 13,14 13 23 14  same as above 48 37'N 122 52'W 48 37'N 122 55'W 48 36'N 122 54'W 48 36'N 122 50'W 48 36'N 122 55'W same as above  T 3 6 N R 2 W Sec. 21 T 3 6 N R 2 W S e c . 17,22 T36N R2W Sec. 17 T36N R2W Sec. 13 T36N R2W Sec. 12 T36N R3W Sec. 1 T 3 6 N R 3 W S e c . 11,14 T37N R3W Sec. 23 T 3 7 N R 3 W S e c . 13,24 T 3 7 N R 3 W S e c . 13 T 3 7 N R 3 W S e c . 1, 12 T37N R2W Sec. 6, 7 T37N R3W Sec. 12 T37N R3WSec. 13  48 36'N 122 57'W 48 36'N 122 59'W 48 37'N 122 59'W 48 36'N 123 01'W same as above 48 38'N 123 00'W 48 37'N 123 02'W 48 41'N 123 02'W same as above same as above 48 4 3 ' N 123 01'W  41  48 4 3 ' N 48 42'N  123 02'W 123 02'W  No. 79 80 81 82 83  Island Waldron Waldron Flattop Spieden Spieden  84 85 86  Spieden Spieden Satellite  87 88 89 90 91 92 93 94  Stuart Stuart Stuart Stuart Sentinel Posey Henry San Juan  95 96 97 98 99  San San San San San  100 101  San Juan San Juan  102 103 104  San Juan San Juan San Juan  105 106 107 108 109 110 111 112 113 114  San San San San San San San San San San  115  San Juan  116  San Juan  117  San Juan  118 119 120  San Juan San Juan San Juan  121  Goose  Juan Juan Juan Juan Juan  Juan Juan Juan Juan Juan Juan Juan Juan Juan Juan  Location Cowlitz Bay Marsh Road to and Sandy Point Area Entire Island Green Point and Dock Area Center of Island around houses and north side of the Island Area around the airstrip West End of the Island Entire Island Co. Rd. to and Turn Point Area Co. Rd., by Schools & Cemetery Above the Reid Harbor Dock Stuart Island State Park Entire Island Entire Island North East End of Island Roche harbor, Resort and Old Rock Quarries Ruben Tarte Co. Park SJINHP, British Camp & Bell Pt. SJINHP, Mt. Young, & Cemetry Old Mitchell Bay Rock Quarry Mitchell Bay Rd, near jet. with West Side Road Cady Mountain Egg Lake Rd. across from Egg Lake Sportsman Lake Euereka Area, Old Kiln Site U.W. Friday Harbor Labs, & Pt. Caution Beaverton Valley Bog and Area Smallpox Bay Trout Lake Lime Kiln State Park Mt. Dallas and Area Mt. Ben, Top and Side Deadman Bay Along the West Side Road McGinitie Property SJINHP, American Camp, Grandma's Cove SJINHP, American Camp, around the Visitor Center SJINHP, American Camp.trail to and around Jakles Lagoon SJINHP, American Camp, Mt. Finlayson & Third Lagoon Cattle Point Picnic Area Mt. Finlayson Swamp Cape San Juan Area, "Goose Control" Entire Island  T.R.S. T37N R3W Sec. 14, 23 T37N R3WSec. 14,15 T36N R3W Sec. 4 T36N R3W Sec. 7 T36N R3W Sec. 6 T36N R4W Sec. 1, 12 T36N R4W Sec. 1 T36N R4W Sec. 2 T 3 7 N R 4 W Sec 21,22, 27, 28 T37N R4W Sec. 20 T37N R4W Sec. 28,29 T37N R4W Sec. 28 T37N R4W Sec. 28 T36N R4W Sec. 2 T36N R4W Sec. 14 T36N R4W Sec. 15 T36N R4W Sec. 23  Lat. & Long. 48 41'N 123 02'W 48 4 2 ' N 123 03'W 48 38'N 123 05'W 48 38'N 123 07'W same as above  T36N T36N T36N T36N T36N  48 36'N 123 06'W 48 35'N 123 07'W same as above 48 35'N 123 10'W 48 34'N 123 10'W  R3W Sec. R4W Sec. R4W Sec. R4W Sec. R4W Sec.  17 26 25 34 35  same as above same as above 4841'N 123 11'W 48 4 1 ' N 123 same as above same as above same as above 48 38'N 123 48 37'N 123 48 36'N 123 48 37'N 123  12'W  10'W 10'W 11'W 07'W  T35N R3W Sec. 6 T36N R3W Sec. 32  48 33'N 48 35'N  123 07'W 123 05'W  T36N R3W Sec. 33 T36N R3W Sec. 34 T35N R3WSec. 1, 12  48 35'N 48 35'N 48 32'N  123 02'W 123 02'W 123 00'W  T35N T35N T35N T35N T35N T35N T35N T35N T35N T34N  48 32'N 123 02'W 48 32'N 123 10'W 48 32N 123 07"W 48 30'N 123 10'W 48 31'N 123 07'W 48 31'N 123 07'W 48 30'N 123 07'W same as above 48 30'N 123 05'W 48 27'N 123 00'W  R3W Sec. 10 R4W Sec. 11 R3W Sec. 18 R4W Sec. 23 R4W Sec. 24 R4W Sec. 24 R4W Sec. 24 R4W Sec. 24,25 R3W Sec. 32 R3W Sec. 11  T34N R3W Sec. 1  48 27'N  123 02'W  T34N R2W Sec. 7  48 27'N  123 00'W  T34N R2W Sec. 7  same as above  T34N R2W Sec. 8 T34N R2W Sec. 8 T34N R2W Sec. 5  48 27'N 48 27'N 48 28'N  122 57'W 122 57'W 122 57'W  T34N R2W Sec. 8  48 27'N  122 57'W  42  No. 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145  Island Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Lopez Turn Yellow McConnel Crane Skull Shaw Shaw Shaw  Location Iceberg Point Agate Beach Co. Park Point Colville Watmough Head Sperry Penn. Camp Nor'Wester Hunter Bay, Bill's Property Jet. of Center Rd. & Mudbay Rd. Shark Reef Sanctuary Center Church Cemetary Rock Point Lopez Hill Hummel Lake Spensor Spit State Park Odlin Co. Park Humphery Head Upright Head. Ferry Terminal Entire Island Entire Island Entire Island Entire Island Entire Island Broken Point, & Yanson Farm Ben Nevis Mt. & Loop Road Neck Point  146 147 148  Shaw Shaw Shaw  149 150 151 152 153 154  Shaw Shaw Shaw Willow Blakely Blakely  155 156 157 158 159  Blakely Blakely Blakely Blakely James  Hicks Bay & Point George U.of W. Caretakers Home site U . of W. Preserve, along Hoffman Rd. Along Smugglers Cove Road San Juan Co. Park Along the Road to Pt. Hudson Entire Island Around the Post Office Blakely Peak & Spencers Cabin Areas Horseshoe Lake Spencer Lake Thatcher Bay Area Road to and S. End of Island Entire Island  T.R.S. T34N R2W Sec. 23,24 T34N R2W Sec. 24 T34N R1W Sec. 21 T34N R l W S e c . 21 T34N R l W S e c . 5 T34N R l W S e c . 6 T34N R2W Sec. 2 T34N R2W Sec. 9 T34N R2W Sec. 3 T35N R2W Sec. 28 T35N R2W Sec. 36 T35N R2W Sec. 23 T35N R1W Sec. 7 T35N R2W Sec. 2 T35N R2W Sec. 1 T36N R2W Sec. 36 T35N R2W Sec. 17 T36N R3W Sec. 23 T36N R3W Sec. 24 T36N R2W Sec. 19 T36N R2W Sec. 5 T 3 6 N R 2 W Sec. 20,21 T36N R2W Sec. 29 T36N R2W Sec. 30 T36N R3W Sec. 25 T35N R2W Sec. 5 T35N R2W Sec. 4 T36N R2W Sec. 33  Lat. & 48 26'N 48 26'N 48 25'N 48 25'N 48 28'N 48 2 8 ' N 48 27'N 48 27'N 48 28'N 48 30'N 48 2 9 ' N 48 3 1 ' N 48 32'N 48 33'N 48 33'N 48 34'N 48 32'N 48 35'N 48 36'N 48 36'N 48 38'N 48 33'N 48 35'N 48 35'N  T36N T36N T36N T35N T36N T36N  R2W Sec. R2W Sec. R2W Sec. RlWSec. RlWSec. RlWSec.  48 48 48 48 48 48  T36N T35N T35N T35N T35N  R1W Sec. 33 R l W S e c . 3,4 RlWSec. 4 R l W S e c . 9, 10 R1W Sec. 14,23  43  27 34 26 9 28 27,34  Long. 122 53'W 122 52'W 122 48'W 122 48'W 122 50'W 122 51'W 122 55'W 122 56'W 122 54'W 122 57'W 122 53'W 122 53'W 122 52'W 122 53'W 122 52'W 122 55'W 122 57'W 123 02'W 123 01'W 123 00'W 122 59'W 122 57'W 122 57'W 123 00'W  48 33'N 122 57'W same as above same as above 35'N 33'N 35'N 32'N 37'N 34'N  122 56'W 122 55'W 122 55'W 122 50'W 122 48'W 122 48'W  48 34'N 122 48'W same as above same as above same as above 48 31'N 122 46'W  Figure 6. Map of collecting locations in the San Juan Islands.  Where possible, collections were made from every habitat and sub-habitat on each island. When accessible, collections were made from the high point on each island. A n attempt to quantify the area covered by collecting was made using an existing grid system. The United States is divided up into a system o f townships each six miles square. Each township is composed o f 36 one mile square sections, thus each island has "x" number o f sections covering the area. Table 8 gives the total number o f sections on each island and the total number o f sections that were visited. The other three columns i n Table 8 reflect the total number o f sub-habitats visited, total number o f mosses collected (collections), and the total number o f species for each island. In some cases, such as Lopez Island, there are 50 sections, but collections were made from only 17 o f them. Several reasons are responsible for not collecting in all o f the sections on an island. In some cases the sections covered mostly ocean and only a tiny piece o f the section was on land, thus not warranting a visit. O n Lopez, there is a large amount o f agricultural land, thus once this habitat had been visited it did not merit repeat trips just to collect i n a specific section. Finally, in some cases, most o f the section was on private land and access was denied. Ecological data were recorded on pre-stamped paper bags, (see Appendix B ). Data recorded included: geographic location, (township, range, section, latitude, longitude), substratum (including vascular plant species i f the moss was an epiphyte), available light, and exposure, (when relevant). Collecting numbers were assigned after the collections were sorted. If, during the identification process, another species was found that was significant enough to be labeled separately, a lower case letter, i.e. a, b, c, was assigned to the original collecting number. Field data were then entered into a Paradox 4.5 database for future analysis. Mosses were placed in herbarium packets and labeled. Voucher specimen are deposited i n the following herbaria: University o f British Columbia, ( U B C ) , University o f Washington, ( W T U ) , U . S . National Park Service, San Juan Islands National Historical Park, (only material collected  45  Table 8. Number o f sections visited on each island.  Island  BLAKELY CLARK CRANE FLATTOP FREEMAN GOOSE HENRY JAMES JONES LITTLE M A C LOPEZ MATIA McCONNELL ORCAS PATOS POSEY SAN JUAN SATELLITE SENTINEL SHAW SKULL SPIEDEN STUART SUCIA TURN WALDRON WILLOW YELLOW  Total No. of Sections  Total No. of Sections Collected In  Total No. of Sub-habitats  No. of Collections  No. of Species  10 1 1 2 1 1 5 2 2 1 50 2 1 88 3 1 75 4 2 16 1 5 9 4 2 11 1 2  9 1 1 1 1 1 1 2 2 1 17 2 1 52 3 1 37 4 2 14 1 4 3 4 2 9 1 2  14 5 11 4 4 2 5 6 9 3 16 9 7 20 7 4 20 6 6 14 3 12 12 9 3 12 3 7  245 53 97 35 17 16 42 43 85 21 570 85 40 2181 78 24 1073 36 90 284 40 132 315 149 29 170 27 40  78 37 51 23 15 10 28 26 47 17 99 47 27 192 46 19 152 28 51 84 25 62 82 55 20 65 19 26  46  on Park Service property). A small study collection was given to the U . S . Fish and Wildlife Service office in Olympia. Identification o f the mosses collected during this study was carried out utilizing the following floras: Lawton (1971), Flowers (1973), Grout (1928-1941), Crum (1984), Crum and Anderson (1981), Frisvoll (1983, 1988), Vitt (1973), plus considerable reference to critically determined herbarium specimens at U B C , especially with reference to taxonomically troublesome taxa.  47  CHAPTER 5 Ecology Both habitat and substrata within it play an important role in the distribution o f mosses. Daubenmire (1968) states that habitat is "usually used to denote a rather specific kind o f living space or environment, i.e. a constellation o f interacting physical and biological factors which provide at least minimal conditions for one organism to live or for a group to appear together." Species diversity is influenced by the complexity o f the habitat, stability, succession, productivity and composition (Slack 1977). Richards (1932) comments, "an obvious approach to the ecological problem is to study exceptional habitats such as caves, hot springs and mountain tops, where the influence o f certain factors is shown in an extreme degree. It has thus happened that we are better informed about the ecology o f the mosses in such places than those of our ordinary woods" Although the unusual habitats in the San Juan Islands were thoroughly explored during the present study, equal attention was also paid to the "common woods", and every attempt was made to gather information on habitat and substratum for each moss collected. The interpretation of that information follows. Moss Species A s Related To Habitat and Sub-habitat The moss flora o f the San Juan Islands is comprised o f 224 species (and varieties) found in a variety o f habitats. For the purpose o f this study the following six general habitat types were established: Maritime, Meadows and Ridges, Disturbed Sites, Rock Outcrops, Wetlands and Woodlands. Because mosses usually occupy microhabitats, these six general habitat types were further divided into 23 smaller vegetation units that were called sub-habitats. (See Table 9 for descriptions and representative locations, and Table 10 for the sub-habitats collected in, on each island, and Appendix C for a complete list o f species associated with each sub-habitat, and see Appendix D for a list o f acronyms).  48  Table 9. Habitats and sub-habitat for the San Juan Islands General Habitat MARITIME  Sub-habitat Code MAR  MSE MCS MEADOWS AND RIDGES  OGS OMP  D I S T U R B E D SITES  ORS CEM DAI DDE ORQ  ROCK OUTCROPS  OOC OON OSN  WETLANDS  WBO WLA WPO WSE WSM WST  WOODLANDS  Description All sites, below 20 feet elevation along the margins of the islands, (excluding seeps),soil banks, rocky shoreline  Example Location all shorelines within the islands  Beach bank seeps, seasonal or continuous, usually of soil or may be rocky Exposed limestone areas along the beach, may be seasonal or continuous wet areas, usually on rock Open grassy slopes usually in full sun, treeless areas without rock outcrops Open meadows, pastures, and lawns, usually in full sun, treeless and often disturbed Open rocky slopes in full sun, grassy areas with scattered small rock outcrops Cemeteries, on old marble, cement, and granite headstones Airstrips, usually mineral soil, active or abandon Developed sites, i.e. housing, commercial buildings, old walkways etc. Old rock quarries, both limestone and sandstone Open calcareous areas, in full sun and dry Open noncalcareous, in full sun, may be basalt or sandstone outcrops Shaded noncalcareous, wet or dry, usually forested sites bogs, sphagnum, carex sedge, or old wet peat deposits, mostly in full sun, but may be shaded  Matia Island, Toe Point Cove  Lakes, along the margins, and on floating logs etc. Ponds, seasonal and continuous wet, may be forested depressions, or in full sun Seeps, non marine, may be forested or non forested, full time or seasonal  San Juan Island, Sportsman Lake  Swamps and marshes, may be seasonal or constantly, wet areas with trees, shrubs or cattails Streams and creeks, mostly constantly wet areas, forested or in full sun  Orcas Island, in the East Sound area  San Juan Island, American Camp  Orcas Island, lawns in Moran State Park Shaw Island, old pastures Orcas Island, Entrance Mountain area  most of the islands most of the islands all of the islands, Stuart Island, Turn Point San Juan Island, Roche Harbor Sucia Island, Fossil Bay area San Juan Island, Lime Kiln Point Orcas Island, top of Mt. Constitution Waldron Island, Pt. Disney Orcas Island, in Moran State Park Orcas Island, Summit Lake San Juan Island, Beaverton Valley area  Orcas Island, in the Mt. Pickett area  Orcas Island, along the Crow Valley road San Juan Island, Egg Lake area Waldron Island, Cowletz Bay Marsh, Lopez, Camp Norwester Orcas island, Cascade Creek  Coniferous forested areas, mixed, Doug Fir. W. Hemlock, Grand Fir, Red Cedar, some Yew and scattered stands of Juniper Deciduous forest areas of Garry Oak, Alder, and Maple  Orcas Island, Moran State Park San Juan Island, Small Pox Bay  MWO  Mixed stands of conifers and deciduous trees, including Madrona  OAO  Old apple orchards, mostly with apple trees, abandoned or active orchards  Orcas Island, Pt. Doughty Sucia Island, trail to Shallow Bay from Fossil Bay Blakely Island, old homestead site  CWO  DWO  49  San Juan Island, English Camp  Table 10. Islands and sub-habitats from which collections were taken. Maritime  ISLAND Blakely Clark Crane Flattop Freeman Goose Henry James Jones Little Mac Lopez Matia McConnell Orcas Patos Posey San Juan Satellite Sentinel Shaw Skull Spieden Stuart Sucia  M A R  X X X X X X X X X X X X X X X X X X X X X X X X  I  M  s  E  I  M C  s  I  o G  s  Meadows & Ridges I  o  M p  X  R  s  I  C E M  I  D A I  I  D D E  X  X  X  I  Outcrops  0 R Q  I  C  X  X  I  o o N  X  X  X X X  X  X  X  X  X  X X X  X  X X  X X X  X  X  X  X X X X X X X  X X X  X  X  X X  X  X X X X  X  X  X  X X X  X  X X X  X  X  X  X  50  I  o s N  X X X  X X X X X  X X  X X X  X X X X X  Willow Yellow  o o  X X  Turn Waldron  o X  X  X  T  Disturbed Sites  X X X X X X X X X X X X X X X X X X  X X X X  X X X X X X  Wetlands I  w B  o  I  w L A  X  I  w p  o  I  w s E  X  I  Woodlands  w s M  X X X  w s T  X  X X X  X  X  X X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X X  I  c w o X X X X  X X X  X  X X  I  X X X X X X X X X X X X X X X X  I  D  I  M  w o  w o  X  X X X  X  X X X  X  X X  X X X X  X X  0 A  o X  X  X X  X  X X X X  X X X  X X  X  X  X  X  X  X  Three general habitats have the highest moss diversity: Woodlands, with a total o f 135 species (60.2% o f the flora), Rock Outcrops with 131 species (58.4%), and Wetlands, with 129 species (57.5%). This high moss diversity is the result o f the availability o f numerous microhabitats and the constant availability o f water. The remaining three general habitats, Disturbed Sites, Maritime, and Meadows and Ridges showed the lowest moss diversity by having fewer than 100 species or 50.0 % o f the flora (Table 11). This lower diversity is probably the result o f less water and fewer sites availability as well as frequency o f "weedy" species (species that occupy sites which have undergone human disturbance and are not usually found elsewhere). Representative species found within each sub-habitat are noted below. The Maritime habitat was divided into the following three sub-habitats; M A R includes all sites below 20 feet elevation along the margins o f the islands, excluding seeps. This subhabitat contained a variety o f substrata and varied from exposed rocky shore lines with beach logs, to humus and duff areas above the beach. A total o f 79 species (35.2 % o f the flora) occurred within this sub-habitat.  Two species, Schistidium maritimum and Ulota  phyllantha,  occurred predominantly on the exposed rocky shores. Above the shoreline, usually on humus and duff, Eurhynchium oreganum, Dicranum scoparium and Plagiomnium insigne, were common. Occasionally, Ulota phyllantha occurred on tree or shrub branches that projected out over the beach bank. Both the M S E (beach bank seeps) and M C S (calcareous beach bank seeps) sub-habitats were characterized by seasonal or continuous water seepage, thus resulting in some unusual microhabitats. The M S E sub-habitat occurred on non-calcareous soil (sometimes on rock or soil over rock) along the beach bank overhang i n full to partial shade. A total o f 28 species (12.5 % of the flora) occurred in these areas. Common species included Amblystegium Eurhynchium praelongum and Pohlia  serpens,  cruda.  The M C S sub-habitat consisted o f soil or soil over rock i n both full shade and full sun, along or just above the shoreline. These calcareous seepages were confined to the few limestone outcrops available on the islands. The limited number o f limestone outcrops determined that 51  Table 11. Moss diversity in the habitats. Legend: M A R , all sites below, 20 ft. elev., excluding seeps, M S E , beach bank seeps, M C S , calcareous beach bank seeps, OGS, open grassy slopes, O M P , open meadows, pastures and lawns, ORS, open rocky slopes, C E M , cemeteries, DAI, airstrips, D D E , developed sites, ORQ, old rock quarries, OOC, open calcareous outcrops, OON, open non-calcareous outcrops, OSN, open shaded non-calcareous outcrops, W B O , bogs, W L A , lakes, WPO, ponds, WSE, seeps, non-maritime, W S M , swamps and marshes, WST, streams and creeks, C W O , coniferous woodlands, DWO, deciduous woodlands, M W O , mixed coniferous and deciduous woodlands, O A O , old apple orchards. Note: Many of the species occur in more than one habitat.  MAR MSE MCS  79 28 11  % Fora 35.2 12.5 4.9  OGS OMP ORS  25 25 53  11.1 11.1 23.6  CEM DAI DDE ORQ  12 21 66 32  5.3 9.3 29.4 14.2  OOC OON OSN  20 115 86  8.9 51.3 38.3  WBO WLA WPO WSE WSM WST  46 56 41 44 51 52  20.5 25.0 18.3 19.6 22.7 23.2  CWO DWO MWO OAO  116 66 88 14  51.7 29.4 39.2 6.25  Sub-Habitat  No. of Species  General Habitat  Total No. of Species  % Flora  Maritime  96  42.8  Meadows & Ridges  73  32.5  Disturbed Sites  74  33.0  Outcrops  131  58.4  Wetlands  129  57.5  Woodlands  135  60.2  52  only 11 species (4.9 %) o f the flora) including Crumia latifolia, Eucladium verticillatum, and Hymenostylium recurvirostre occurred within this sub-habitat. Tufa formations were often formed within these sites when the water supply was constant. The Meadows and Ridges habitat was divided into the following three sub-habitats: open grassy slopes (OGS), open rocky slopes (ORS), and open meadows and pastures ( O M P ) . Open grassy slopes ( O G S ) are usually treeless, covered by grasses, and i n full sun. Common substrata included dry soil, limited humus and duff, small rocks and occasionally pieces o f decayed wood. A total o f 25 species (11.1 % o f the flora) occurred within this sub-habitat. Common species included Brachythecium albicans, Polytrichum juniperinum,  and Weissia controversa.  The open  rocky slope (ORS) sub-habitat consisted o f small, l o w rock areas scattered throughout the grassy slopes. Species on these low rocky areas included Polytrichum piliferum,  Racomitrium  ericoides, Didymodon vinealis, and, occasionally, Hedwigia stellata. A total o f 53 species (23.6 % o f the flora) occurred within this sub-habitat. Finally, the O M P or open meadows and pastures sub-habitat consisted o f pastures, lawns or open meadows in full sun. These areas were often disturbed and usually had more water available to them than the other two sub-habitats. Only 25 species (11.1 % o f the flora) were found within this area, usually on damp or dry soil. Two species, Pseudoscleropodium  purum and Rhytidiadelphus squarrosus, occurred only in this sub-  habitat. Other common species included Brachythecium albicans, Bryum pseudotriquetrum,  and  Leptodictyum riparium. In spite o f the availability o f extra moisture this sub-habitat was fairly low in species diversity, possibly the result o f disturbance and competition from vascular plants. The Disturbed Sites habitat was divided into four sub-habitats based on the type of disturbance that had occured. Sometimes, imported headstones in cemeteries bring in an "exotic" moss species; thus, cemeteries ( C E M ) on each o f the islands were checked for this possibility. Only 12 species (5.3 % o f the flora) were found in this sub-habitat in spite o f headstones constructed from a variety of rock materials, and no "exotic" introduced species were found. Common species found on the headstones included Dicranoweisia  53  cirrata, Didymodon  vinealis,  Homalothecium pinnatifidum Brachythecium  and Tortula muralis.  albicans and Ceratodon  Species found on dry soil included  purpureus.  Airstrips (DAI) both active and abandoned, were often composed o f raw mineral soil. A total o f 21 species (9.3 % o f the flora) occurred within this sub-habitat. C o m m o n species included Polytrichum juniperinum,  Racomitrium heterostichum and Ceratodon  purpureus.  Several uncommon species found occasionally in or along the edges o f the tire ruts were Physcomitrium pyriforme and Pleuridium acuminatum.  Developed Sites ( D D E ) included old  walkways, and housing or commercial building sites. These disturbed areas were usually colonized by "weedy" species that often did not occur in other less disturbed sub-habitats. A total o f 66 species (29.4 % o f the flora) was found in this sub-habitat. The following species were usually present in these sites: Tortula muralis, Grimmia pulvinata, and Ceratodon  purpureus.  O l d Rock Quarries ( O R Q ) were common on several o f the islands, and were composed o f limestone or sandstone. Thirty-two species (14.2 % o f the flora) were found within this subhabitat and the most common substratum was rock. Occasionally, trees grow in some o f the old quarries, thus providing both shade and additional microhabitats for several species. These old quarries are usually open pits exposed to full sun and thus were very dry sites. Common species on rock included Didymodon vinealis, Racomitrium ericoides, and Ptychomitrium  gardneri.  The general Rock Outcrop habitat was divided into three sub-habitats, based on the amount o f light exposure and the composition o f the rock material. O O C , or open calcareous rock outcrops, were dry sites in full sun and usually composed o f limestone. Only 20 species (8.9 % of the flora) occurred within this sub-habitat and included the following species: capillare, Tortella tortuosa, and Gymnostomum aeruginosum.  Bryum  O O N or open non-calcareous  rock outcrops were large open areas containing a variety o f rock materials. These large exposed areas provided numerous microhabitats and 114 species (50. 8 % o f the flora) occurred within this sub-habitat. The following species were found: Pseudobraunia stellata, Isothecium cristatum, and a variety o f species o f Racomitium.  54  californica,  Hedwigia  When the crevices or  rock ledges provided shade or occurred on the north slope, the following species were usually present: Anacolia menziesii, Amphidium californicum, Claopodium  bolanderi.  Shaded non-calcareous outcrops ( O S N ) were located within the forested areas, thus the shade helped to increase the available moisture in the microhabitats. A total o f 87 species (38.8 % of the flora) was collected from this sub-habitat. Common species included Amphidum californicum and A. lapponicum, Bartramiapomiformis, menziesii.  Encalypta ciliata, and Anacolia  When moisture was present as seepage the following species were often present:  Porotrichum  bigelovii, and Dichodontium  pellucidum.  Although several o f the islands have fresh water lakes, for the most part there is not an abundance o f surface water present. In many cases, where a stream or lake did exist it has been modified. When a fresh water source is present (disturbed or undisturbed) it forms a distinct microhabitat that may contain a variety o f species that are dependent on the high moisture conditions. Therefore the Wetlands habitat was divided into five sub-habitats based on the type or source o f the water available. The highest moss diversity occurred in the lake ( W L A ) subhabitat, with 56 species (25 % o f the flora). This sub-habitat contained a wide variety of substrata including floating logs (or docks), over-hanging tree or shrub branches, standing dead trees, and the shoreline which varied from rock cliff faces to humus and duff banks. This variety of substrata probably is responsible for the high moss diversity o f this area. A few o f the species associated with the wetter substrata included Calliergon giganteum, Climacium dendroides and Sanionia  uncinata.  The Streams ( W S T ) sub-habitat contained 52 species (23.2. % o f the flora). This subhabitat was best represented on Orcas Island where several large creeks or streams drain from Mt. Constitution. Species associated with these streams included Scleropodium Brachythecium frigidum,  and Fissidens ventricosus.  obtusifolium,  In one area where the creek flowed over a  small lens o f limestone Fissidens grandifrons, a calcicole, appeared i n several locations. Because this sub-habitat also included the bounding creek or stream bank, it increased the number o f substrata, thereby increasing the moss diversity. The Swamp and Marsh ( W S M ) sub55  habitat showed the third highest moss diversity with 51 species (22.7 % o f the flora). This high diversity again is the result o f the large number o f substrata occurring in the sub-habitat. These seasonal or constantly wet areas were associated with either trees, shrubs or cattails. Areas dominated by cattails or covered by dense shrub layers generally were not species rich. This is probably a response to the vascular plants that out-compete the mosses by reducing the available sites for colonization. O n the other hand, when trees were present, the moss diversity increased as a result of the increased number o f available microhabitats. Species commonly associated with this sub-habitat included Amblystegium serpens and Drepanocladus aduncus, both found on wet soil, Onocophorus wahlenbergii on the base o f trees, and Plagiomnium medium on decayed wood. The B o g and bog like ( W B O ) sub-habitat was one o f the most unexpected i n the islands. It was restricted to San Juan and Orcas Islands. The bogs were composed o f either sedges (Carex sp.), sphagnum, or were old peat deposits, and in many cases they had experienced some form of disturbance. A total o f 46 species (20.5 % o f the flora) occurred in these bogs. Dominant species included a variety o f species of Sphagnum and Aulacomnium palustre.  Pohlia  sphagnicola and Bryum uliginosum were found only in the Orcas Island Sphagnum bogs. Forty four species (19.6 %) were found in the non-maritime seep sub-habitat (WSE). This low number o f species is most likely due to the limited amount o f fresh water sites within the islands. The wetland with the lowest moss diversity was the Pond ( W P O ) sub-habitat. Most o f the man- made ponds were extremely poor in species composition because o f the continuity o f disturbance, either from grazing animals or fluctuating water levels in response to irrigation needs. The seasonal or constantly wet forest floor depressions that ponded water during wet periods had a high species composition and thus are responsible for most o f the 41 species (18.3 %) found in this sub-habitat. Common species in the forested sites included  Fontinalis  antipyretica and Leptodicytum riparium. Drepanocladus crassicostatus and Bryum flaccidum were found in a shallow depression on wet soil along a road in full sun.  56  The Woodlands habitat was divided into four sub-habitats based on tree species composition. While there are areas on M t . Constitution that have a few old trees, true old growth forest is not represented within the islands. Thus the coniferous woodlands ( C W O ) are all second growth Pseudotsuga menziesii (Douglas fir), Tsuga heterophylla (western hemlock), Abies grandis (grand fir), and Thuja plicata (red cedar). Taxus brevifolia (yew) is also scattered throughout the area although it is not abundant, and Juniperus scopulorum (Rocky Mountain juniper) occurs in small clusters on some o f the drier sites. This sub-habitat had the highest diversity with a total o f 116 species (51.7 % o f the flora). A wide range o f substrata were found throughout the sub-habitat and included: humus and duff, damp and dry soil, decayed wood, epiphytes, and a variety o f small rock types. Forest floor species composition is usually rather high as there is less litter accumulation. Common forest floor species included robusta, Trachybryum megaptilum, and Pleurozium schreberi. trichophylla,  Racomitrium lawtonae, and Cynodontium jenneri.  Rhytidiopsis  Rock species included Dicranella  Grimmia  heteromalla and  Pohlia nutans often occurred on soil banks along the trails. Species associated with the coniferous tree trunks included Hypnum circinale and Dicranum tauricum.  W i t h the exception  o f one juniper tree on Posey Island which was leaning, mosses were not found on this substratum. The deciduous woodlands ( D W O ) sub-habitat is composed o f stands o f Garry oak, alder, or big leaf maple trees. This sub-habitat is scattered throughout the islands but has been greatly impacted by human development. The predominant substratum found in this sub-habitat are the tree trunks and branches which provide excellent sites for a variety o f corticolous species. Common species found on the trunks o f trees include Plagiomnium venustum, Claopodium crispifolium,  Homalothecium nuttallii, Isothecium myosuroides, and Orthotrichum  Species on branches included Tortula ruralis, Orthotrichum consimile, and Ulota  lyellii. obtusiuscola.  A total o f 66 species (29.4 % o f the flora) occurred i n this sub-habitat. These tree stands are often isolated patches in the middle o f large open areas where grasses dominate the ground cover.  57  This fact, combined with the accumulation o f leaf litter, acted to prevent an abundance of moss species from establishing under these stands o f trees. M i x e d Woodlands ( M W O ) were composed o f both coniferous and deciduous trees as well as madrona. This sub-habitat had 90 species (40.1 % o f the flora). These wooded areas often had an understorey o f Gaultheria shallon (salal) or Holodiscus discolor (ocean spray) thus decreasing the availability o f ground sites. The deciduous trees consistently had the same epiphytic species mentioned above, while the coniferous trees usually had Hypnum circinale and Dicranum tauricum on the trunks. Arbutus menziesii (madrona) rarely had epiphytic species because o f the shedding nature o f the bark. Occasionally, Eurhynchium oreganum or Isothecium myosuroides could be found growing up the base o f an old, well established madrona tree. O l d Apple Orchards ( O A O ) had the lowest moss diversity for woodlands, with only 14 species (6.2 % o f the flora). This is because the only substratum available for colonization was on the tree trunk or branches. Usually these orchards were in open grassy areas and thus ground species could not become established. Occasionally in the abandoned orchards fallen branches or trunks would provide additional substratum for such species as Aulacomnium  androgynum.  The  epiphytic species on trees and branches were consistently the same as in the deciduous tree subhabitat. Since mosses are "well known to be sensitive to subtle differences in micro-environment" (Bates 1982), substratum plays an important role in their distribution.  58  Moss Species A s Related To Substratum Classification o f bryophytes on the basis o f substratum has been carried on since the late 1800's (Gams 1932). While "most bryophytes have a sharply defined and rather narrow ecological range" (Richards 1932), others occur on a variety o f substrata. Knowing whether a moss has an obligate or facultative relationship with it's substratum is often important when making a species determination. For this study, 18 substratum categories were established and data were recorded for each species collected. See Table 12. for these categories and their definitions. The rock substratum showed the highest moss diversity with 160 species (71.4 %) of the flora. This probably results from the vast number o f microhabitats and the variety of rock material available for colonization. A n additional reason for this large number o f species is that most rock outcrops do not accumulate much soil: therefore, vascular plants can not become well established, and the competition for the bryophytes is lessened. The second highest diversity occurred on the soil substratum, with 141 species (62.9 % o f the flora). The lowest diversity occurred in the aquatic substratum, with only 15 species (6.6 % o f the flora). This reflects the lack o f abundance o f fresh water within the islands, thus fewer aquatic species can become established. Peatlands showed the second lowest moss diversity, with 28 species (12.5 % o f the flora), again emphasizing the lack o f suitable sites. Living-wood, dead-wood and humus and duff all represent less than 50.0 % o f the flora. See Table 13. for a complete summary and Appendix E for a list o f species associated with substratum. Rocks offer a wide variety o f microhabitats for mosses to colonize. These sites include fissures and crevices, ledges, smooth or rough surface texture, and small "caves". Mosses growing on rock may be either obligate or facultative epiliths (Gams 1932). Several o f the obligate species found during this study were Andreaea  megistospora,  Grimmia trichophylla, Orthotrichum rupestre and Schistidium maritimum. Facultative species found on this substratum included Bryum capillare, Eurhynchium  59  praelongum,  Table 12. Definitions of substrata. General Substratum Water  Substratum (sub-code)  Definitions  AQ  Aquatic  BL DW FL  Beach logs Decayed wood Floating logs  Living-Wood  EP FO  Epiphytic Folicolous  Humus & D u f f  HD  Humus and duff  Peat  PE  Peatlands  RA RB RC RG RL RO RS  Slate Basalt Conglomerate Granite or acidic Calcareous & cement U n k n o w n material Sandstone  SC SD SW  Clay Dry Wet  Dead-Wood  Rock  Soil  60  Table 13. Moss diversity according to substratum. Legend: The percentages in this table reflect the occurrence of a species on more than one type of substratum. A Q , aquatic, B L , beach logs, D W , decayed wood, EP, epiphytic, FL, floating logs, H D , humus & duff, PE, peatlands, RA, slate rock, R B , basalt rock, RC, conglomerate rock, R G , granitic, acidic rock, R L , cement and calcareous rock, RO, unknown rock, RS, sandstone rock, SC, clay soil, SD, dry soil, SW, wet soil. Substratum  %  %  Combined Substratum Aquatic  Total No. of Species 9  4.0  Dead-wood  82  36.6  29.4 0.44  Living-wood  66  29.4  55  24.5  Humus & Duff  56  25.0  PE  28  12.5  Peat  28  12.5  RA RB RC RG RL RO RS  5 17 48 26 76 149 25  2.2 7.5 21.4 11.6 33.9 66.5 11.1  Rock  163  72.7  SC SD SW  33 109 74  14/7 48.6 33.0  Soil  142  63.3  AQ  No. of Species 9  4.0  BL DW FL  4 79 1  1.78 35.2 0.44  EP FO  66 1  HD  61  Hypnum cupressiforme, Isothecium myousorides, Tortula princeps, and T. ruralis. These species are found on rock because a shallow layer o f humus has accumulated on the rock surface, thus allowing them to become established. The chemical nature o f rock also plays an important role i n determining which species w i l l be able to survive on a particular substratum. Often the p H o f a substratum can be estimated on the basis of the moss species found growing on it. This is particularly true with the obligate calcicoles such as Crumia latifolia, verticillatum,  and Gymnostomum aeruginosum.  Andreaea rupestris, Bartramiapomiformis, Schistidium  Eucladium  Siliceous or acidic rock obligates include  Encalypta ciliata, Pohlia cruda, and  apocarpum.  Six types o f rock material were easily distinguished in the field; when the nature of the substratum was known, it was recorded. Unfortunately much o f the time the rock material was not easily identified: therefore, 146 species, or 65.1 % o f the flora, were recorded as "rock unknown" (RO). A brief discussion o f each rock substratum follows. Since slate ( R A ) was not a common rock material within the islands, only 5 species (2.2 % o f the flora), occurred on this substratum. T w o species found on this substratum were Amphidium lapponicum and Schistidium  maritimum.  Basalt ( R B ) was also an uncommon rock substratum; only 17 species (7.5 % o f the flora), were found on this substratum. Representative species included Bryum dichotomum, Pseudobraunia californica,  and Schistidium maritimum.  Conglomerate  rock (RC) formations were on a restricted number o f the islands, but the rough texture o f this type o f rock material plus the diversity o f rock fragments in it provided habitat for 48 species (21.4 % o f the flora). C o m m o n species on this substratum included Claopodium bolanderi, Didymodon vinealis var. vinealis, Grimmia trichophylla, and Hypnum subimponens. Granite rock material (RG) was restricted to glacial erratic boulders that were scattered throughout the islands. Crum (1973) comments in a discussion on the moss 62  flora of the Great Lakes area, "glacial erratics present some variety in substrate, o f slight importance"; thus these erratic boulders often have unique species associated with them. This is especially true when the rock material differs substantially from the surrounding surface material. Therefore, these rocks were o f particular interest. Unfortunately they did not show this unique pattern o f unusual species and all o f the species found on them were common to the islands in general. A m o n g the species found were  Dicranoweisia  cirrata, Orthotrichum rupestre, Ptychomitrium gardneri, and Ulota phyllantha, all species that prefer an acidic substratum. A total o f 26 species (11.6 % o f the flora), occurred on these erratic boulders. Although sandstone (RS) occurred on several o f the islands, only 25 species, (11.1 % of the flora) were on this substratum. The low species diversity is probably the result of the unstable surface o f the sandstone which makes establishment difficult. Most o f the species on this substratum were in rock crevices or fissures. The following facultative sandstone species included Antitrichia Leucolepis acanthoneuron.  californica,  Eurhynchium oreganum, and  Obligate rock species included: Dicranoweisia  Didymodon vinealis var. vinealis, and Racomitrium  cirrrata,  ericoides.  Species composition varied according to the type o f calcareous rock ( R L ) material, and the presence or absence of water on the substratum. Since cement or concrete is composed o f lime it was included in this substratum group. These cement structures included walls, curbs, chimneys, and dams. Species associated with dams often reflected the additional moisture that was present, and included the following species Didymodon tophaceus, Hymenostylium recurvirostre, and Gymnostomum aeruginosum.  Species associated with drier sites such as curbs and walls included:  Didymodon vinealis var. vinealis, D. vinealis vat.flaccidus,  and Tortula muralis.  Naturally occurring limestone deposits in conjunction with seepages contained very interesting species. There, obligate calcicoles such as Crumia latifolia, verticillatum,  Eucladium  Gymnostomeum aeruginosum and Hymenostylium recurvirostre all could 63  be found, occasionally forming tufa deposits. Dry limestone outcrops on the other hand often had the same species as found on cement, along with the following species: Didymodon fallax, Orthotrichum anomalum, and Schistidium apocarpum.  When both the  wet and dry limestone substrata are combined, a total o f 76 species (33.9 % o f the flora) are represented. The terrestrial substratum was divided into the following three general categories: humus and duff (HD), peatlands (PE), and soil. Soils were further divided into soil dry (SD), soil wet (SW), and clay soil (SC). These divisions were based on the moisture content, texture and amount o f organic material present at each o f the collecting sites. In general the soils within the islands are composed o f glacial till and outwash, are well drained, and lack large amounts o f loam. Thus, they are not very rich soils and usually have been exposed to recent disturbance. Since moss diversity is dependent upon soil richness, moisture, shading and the amount o f disturbance present, it was unexpected to find 141 species (62.9 % o f the flora) occurring on this substratum. A n examination of the species composition makes this large number easier to understand. W i t h the exception o f a few species, most of the mosses found were those that colonize disturbed areas. The largest number, 109 species (48.6 % o f the flora), occurred on dry soil (SD) and included the following species: Bryum capillare, Ceratodon purpureus, hygrometrica, and Polytrichum juniperinum,  Funaria  all i n sunny, open areas, often mixed with  the surrounding vascular plant weeds and grass. Atrichum selwynii and  Dicranella  heteromalla were found i n shaded sites on bare mineral soil along trails and on upturned tree root systems. Most o f the above species respond well to soil disturbance and thus were found on over 50 % o f the islands. Wet soils (SW) occurred along stream banks, lake margins and in seepy areas. Common species found in these locations included Brachythecium frigidum, Bryum pseudotriquetrum,  Colliergonella  cuspidata, and occasionally, Philonotis fontana.  A  total o f 74 species (33.0 % o f the flora) occurred on this substratum. Although this l o w 64  number o f species does not reflect the normal species richness often found in these moist soil sites, the lack o f suitable wet soil habitats limited the potential for a large number o f species to become established. A s a result o f the limited number o f clay deposits known from the islands, only 33 species (14.7 % o f the flora) occurred on this substratum. Although the clay soil (SC) substratum was restricted, the following two species (that are confined there to clay deposits) were collected: Dicranella  howei, and Dicranella pacifica.  Other species found  on, but not restricted to, the clay substratum included: Fissidens bryoides, limbatus, Leptobryum pyriforme, and Pleuridium  Fissidens  acuminatum.  Humus and duff (HD) is described as the organic constituent o f the soil, and is composed o f partially decomposed plant and animal litter (Lincoln et al. 1982). Thus, "humus is a factor o f first importance" (Grebe 1917), and often provides a suitable substratum over soil and rock for the establishment o f mosses that otherwise would not be present. A total o f 55 species (24.5 % o f the flora) occurred predominately in the coniferous woodlands. A t the higher elevations in the M t . Constitution area, extensive mats o f Rhytidiopsis robusta, Trachybryum megapitlum, and Rhytidiadelphus were found. Lower elevation species included Dicranum scoparium,  triquetrus  Eurhynchium  oreganum, Plagiomnium insigne, and occasionally Hylocomium splendens. Thin layers of humus often form in more open areas over rocks allowing the following species to become well established: Tortulaprinceps,  Tortula ruralis, and Isothecium  myosuroides.  Peatlands (PE) generally are low in species diversity especially when o f small size, and usually species o f Sphagnum dominate this substratum (Schofield 1976). Although low in diversity (only 28 species, or 12.5 % o f the flora), these acid-rich substrata contained some o f the more unusual species in the islands. T w o species, Bryum uliginosum and Pohlia sphagnicola, were found only within the bog complex. Common Sphagnum species included S. fuscum, S. recurvum and S. squarrosum.  65  A m o n g the  other common species found were Aulacomnium palustre, Pleurozium schreberi, and Polytrichum  strictum.  Aquatic ( A Q ) bryophytes occur either as floating mats, or attached to rocks or wood fragments in streams and lakes. The following species often formed floating mats: Drepanocladus  aduncus, Fontinalis antipyretica, and Leptodictyum riparium.  Species  often attached to rocks i n or along the margin o f lakes, i n streams, or i n waterfalls include Brachythecium frigidum, Fissidens ventricosus, and Porotrichum bigelovii.  Fissidens  grandifrons, a calcicole, occured only in those streams where enough calcium was being leached into the water from the surrounding limestone. A total o f 15 speceis (6.6% o f the flora) was found in aquatic areas. Floating logs (FL), decayed wood ( D W ) , and beach logs ( B L ) were combined as dead-wood substrata. Together, 86 species (38.3 % o f the flora) occurred on these substrata. Decayed wood, which included rotten logs, stumps, and standing dead trees, had the highest diversity with 76 species (33.9 % o f the flora). Common species found on this substratum included Dicranum fuscescens, Rhizomnium glabrescens, and Tetraphis pellucida.  Beach logs ( B L ) provided an extremely poor substratum because o f  their proximity to salt water and instability. Thus only 4 species (1.78 % o f the flora) occurred on these logs. Bryum capillare and Ulota phyllantha were common, while Orthotrichum lyellii and Schistidium maritimum were found only once. Floating logs (FL) usually half submerged, were found in several o f the lakes. A total o f 6 species (2.6 % of the flora) was found on this extremely wet substratum. The following species were the most common: Bryum pseudotriquetrum  and Philonotis  fontana.  Species occurring on living-wood were divided into two substratum groups: epiphytic (EP), including those species growing on the trunks or branches o f trees or shrubs, and foliicolous (FO), including those species growing on the leaves o f trees or shrubs. Vitt et al. (1973) reported the first occurrence o f this substratum for Western North America when four species o f Orthotrichum were found on Thuja plicata leafy 66  branches. Only one species, Orthotrichum consimile, was found on Thuja plicata leafy branches confined to a shallow damp drainage on Orcas Island. This tree was within a small grove of Acer macrophyllum trees which probably served as the dispersal source for this Orthotrichum.  Thus, the close proximity to a good source o f spores, and the  increased humidity from the drainage, combined to allow this Orthotrichum to colonize the Thuja. The epiphytic (EP) substratum has both obligate and facultative species occurring on usually deciduous trees. Obligate mosses included Orthotrichum consimile, O. lyellii, Viola megalospora, and Ulota obtusiuscula. rubra (red alder), Acer macrophyllum  A l l occur on the limbs or trunks of Alnus  (bigleaf maple), and Quercus garryana (Garry  oak). Occasionally, these species o f moss were found on Holodiscus discolor (ocean spray). Isothecium cristatum and Scleropodium  cespitans were usually found near the  base o f deciduous trees. A few o f the facultative species included Antitrichia curtipendula, Homalothecium fulgescens, Isothecium myousroides,  Metaneckera  menzesii, Neckera douglasii and Zygodon viridisimus var. rupestris. These facultative epiphytes occurred on both tree trunks and on rocks. A total o f 66 species (29.4 % o f the flora) was found on this substratum. Since a geologic formation or unit is composed o f a variety o f different types o f rock substrata, data were collected to determine i f there was any correlation between these units and the distribution o f mosses.  67  Moss Species A s Related To Geologic Unit While numerous studies have been made concerning the relationship o f mosses to rock substrata (Nagano 1969), it appears that little research has been carried out that relates entire geologic units to species diversity. Therefore, during this study, the geologic units from which any moss was collected were recorded. See Table 14. for a complete summary. Two formations, the Garrison Terrane and Haro Formation were discussed in the geological section o f this paper yet collections were not made from either o f these formations because access was denied or the formation was too small to easily identify i n the field. See Appendix F for a complete list o f species associated with geological unit. The Constitution Formations (CF) showed the highest diversity, with 188 species 83.9 % o f the flora) occurring on this geologic unit. This high species diversity may result from several factors. First, the Constitution Formation occurs widely among the islands, covering all o f Crane and Turn, and most o f San Juan, and Shaw. O n Orcas Island it covers several extensive areas, including the M t . Constitution and M t . Woolard regions, (see Figure 3). Thus, it is one o f the largest geologic units (38.5 % o f the area) represented in the islands. Second, most o f the higher elevation areas that Deither et al. (1995) proposed to have been above water and ice free by 13,500 yr. B P occur within this formation. Thus, it is possible that these areas, having been available for occupation earlier, were colonized by more species. Finally, at the present time, these higher areas tend to be characterized by greater precipitation than other elevations in the islands and, thus, increase the potential for a variety of species to colonize or presist in the area. Two other geologic units contained over 50 % o f the flora. They were the Turtleback Complex (TC) with 151 species (67.4 % o f the flora), and the East Sound Group (EG) with 121 species (54.0 % o f the flora). Both o f these geologic units occur predominately on Orcas Island where they form large bands o f material along the northern side o f the island. Jones Island is also composed entirely o f the East Sound Group. Although the East Sound Group represents only 5.28 % o f the area within the islands, the variety o f rock material found in  68  Table 14. Moss diversity as related to geological unit. Legend: CF, Constitution Formation, C K , Chuckanut Formation, D B , Deadman Bay Volcanic, E G , East Sound Group, F C , Fiadalgo Complex, LF, Lummi Formation, L S , Lopez Structural Complex, N G , Nanaimo Group, OC, Orcas Chert, QC, Quaternary Cover, SG, Spieden Group, T C , Turtleback Complex. Geological Unit  N o . of Species  %  % o f Flora  o f Island A r e a CF  38.54  188  83.9  CK  .96  73  32.5  DB  .63  60  26.7  EG  5.28  121  54.0  FC  10.06  96  42.8  LF  3.81  60  26.7  LS  1.87  65  29.0  NG  4.84  98  43.7  OC  12.61  104  46.4  QC  11.72  74  33.0  SG  .45  68  30.3  TC  11.21  151  67.4  69  this formation substantially increased the species diversity. The high moss diversity found on the Turtleback Complex probably is the result o f both the variety o f rock material that occurs in this formation, and the fact that this complex also appears at the higher elevations on Orcas Island. Two species, Andreaea megistospora and A. rupestris,  were restricted to the  Constitution Formation. A. megistospora was found only in the M t . Constitution area on Orcas Island. Andreaea rupestris also occurred on M t . Constitution, but a historical collection from the M t . Dallas area on San Juan Island indicates that, at one time, it occurred on both of the islands. The following geologic units had between 40 and 50 % o f the flora: Orcas Chert (OC) with 104 species (46.4 % o f the flora), the Nanaimo Group ( N G ) with 98 species (43.3 % of the flora), and finally the Fidalgo Complex (FC) with 96 species (42.8 % o f the flora). While these three units were limited in distribution they contained some o f the more unusual rock materials with respect to the geology o f the islands. Most o f the limestone deposits occurred i n the Orcas Chert, thus the obligate calcicole species were found on this unit. The Nanaimo Group, composed of sandstone, conglomerate, and shale, and the altered igneous rocks rich in greenschist found in the Fidalgo Complex, all provided a variety o f substrata for the colonization o f mosses. Thus, the greater diversity o f substrata probably compensated for the limited distribution o f these geologic units. Less than 40 % o f the flora occurred on the Chuckanut Formation ( C K ) and Quaternary Cover (QC). The Chuckanut Formation, with 73 species (32.5 % o f the flora), is found, in the islands, only on Matia, Patos and Sucia Islands. It is composed o f non-marine sandstone (a poor substratum for bryophytes), and conglomerate, which provided a variety o f substrata for colonization. While the Quaternary Cover occurs on over half o f Lopez and Waldron Islands, it is composed o f glacial tills and sediments. Thus, it is reasonable to assume that the 74 species (33.0 % o f the flora) were found on substrata other than soils.  70  Relatively low moss diversity occurred on the L u m m i Formation (LF) and Deadman Bay Volcanics ( D B ) with only 60 species (26.7 % o f the flora), the Lopez Structural Complex (LS), with 65 species (29.0 % o f the flora), and the Speiden Group (SG), with 68 species (30.3 % o f the flora). With the exception o f the Spieden Group, which occurs only on Spieden Island, these units occur mainly on Lopez and Orcas Islands. Thus, their restricted distribution may determine the low moss diversity. Other than the calcicole species there seems to be a weak correlation between species distribution and geologic unit. It also seems reasonable that the higher moss diversity occurs on the largest geologic unit, probably as a result o f the increased number o f substrata for colonization. Thus, while understanding the geologic formations is important in helping to locate good collecting sites, there is no strong correlation between species distributions and the geologic units. Two unusual features within the San Juan Islands merit particular attention, these are the limestone deposits (mentioned above) that occur scattered throughout the islands, and the bogs and bog like areas found on San Juan and Orcas Islands.  71  Unusual Habitats A n d Substrata Limestone Deposits One o f the most interesting features within the San Juan Islands is the numerous limestone deposits. Danner (1966) surveyed the islands for these deposits and developed a list o f 107 sites. The majority are dry and have been extensively disturbed by quarrying over the years. Although these dry, limestone sites host the calcicolous species Didymodon fallax, Orthotrichum anomalum, and Schistidium apocarpum, for the most part they are l o w in species diversity. When the limestone deposits occur in conjunction with freshwater (i.e. a stream, spring, or seepage) then a distinctive group o f calcicole species may be present, and tufa deposits are often formed. A m o n g the species found in these wet areas are Crumia latifolia, Eucladium verticillatum, aeruginosum, and Hymenostylium recurvirostre.  Gymnostomum  Several o f these species were involved  in extensive tufa formation. Tufa is a "variety o f calcium carbonate ( C a C O ) deposited i n freshwater" (Danner 1966), and according to Flugel (1982) it is often "...formed by the precipitation o f calcite onto plants, (usually algae and mosses)...". Parihar and Pant (1975), comment that the dense moss cushions provide "spongy surfaces which can absorb, retain and expose copious thin films o f water for effective evaporation and consequent diffusion o f C O from the calcareous spring water thus causing the precipitation o f C a C O (calcite)". These tufa formations occur only i n areas where the water carries a high percentage o f carbonates ( C O ). Shacklette (1965) investigated a stream i n Alaska containing 8.8 % carbonates and found only bryophytes growing i n and around the stream. H e further commented that "the mosses were so heavily coated with calcite that only the tips were pliable". Because the growth rate o f mosses exceeds the rate o f carbonate deposition "the process o f getting cemented below and growing above is continued and the tufa also "grows" up" (Parihar and Pant 1975). Crum (1973) names these tufa formations based on species composition; thus, a tufa composed o f Eucladium verticillatum would be called a  72  eucladiolith, i f it were composed o f Didymodon tophaceous it would be a didymodonolith. Several o f these tufa deposits were noted on the islands, one composed mostly of Crumia latifolia and another o f Eucladium verticillatum.  Other sites high in  carbonates did not have tufa formations but still supported the following species: Didymodon tophaceous, Gymnostomum aeruginosum, and Hymenostylium  recurvirostre.  Although there is a substantial amount o f literature concerning tufa deposits (Crum 1973, E m i g 1918, Parihar & Pant 1975, Pentecost 1996, Richards 1932, Richardson 1981, Shacklette 1965, and Taylor 1919), none deals with moss tufa deposits in Washington State. Although Danner (1966) discusses and maps the tufa deposits, he states that "Algae and mosses also are able to induce the precipitation o f calcium carbonate in the form o f tufa, but their contribution to the formation o f deposits in western Washington has not been investigated". The tufa deposits in the San Juan Islands represent one o f the more unusual microhabitats within the state o f Washington. U p until the summer o f 1996 (when it was found on the Olympic Peninsula), Crumia latifolia was known in Washington from only the San Juan Islands. Therefore, additional field work throughout the state is needed to provide a better understanding o f the distribution patterns for these unusual calcicole mosses. Finally, because "...limestone is one o f the most commercially important rocks..." (Danner 1966), it would be prudent to locate and protect these unusual calcareous microhabitats before they have been exploited commercially. Bogs and B o g Like Habitats On the other end o f the p H spectrum, peatlands also provide some o f the more unusual habitats i n the islands. C r u m (1988) defines a bog as "an ombrotrophic peatland, that is, one deriving water and nutrients only from the atmosphere; a highly acid and nutrient-poor peatland dominated by Sphagnum".  These water-logged environments  provide a very distinct substratum for vascular plants and mosses. Typical vascular plants confined to the "bogs" include Drosera rotundifolia (sundew), Eriophorum 73  chamissonis,  and E. gracile (cotton-grass), Kalmia microphylla (western swamp laurel), Ledum groenlandicum  (Labrador-tea), Pinus contorta (lodgepole pine), and two species o f  Vaccinium, V. oxycoccus var. intermedium (wild cranberry) and V. uliginosum var. occidentale (bog billberry). Mosses include Sphagnum capillifolium, henryense, S. magellanicum, S. palustre, subsecundum, Aulacomnium  S. fuscum, S.  S. recurvum, S. rubellum, S. squarrosum, S.  androgynum, Bryum uliginosum, Colliergonella  Pleurozium schreberi, Pohlia sphagnicola, and Polytrichum  cuspidata,  strictum.  In 1934, Rigg and Richardson discussed the development o f eight sphagnum bogs in the San Juan Islands. Peat cores made during their study indicated that these bogs formed in post-glacial depressions, that formerly were lakes, and with the exception o f Blakely bog (which has a sand bottom), all o f the peat deposits rest on the typical blue clay found throughout the region. A survey o f Washington State peat resources, published by Rigg's in 1958, listed all o f the known peat deposits and sphagnum bogs within the islands, and gave detailed location information. In two publications (Rigg and Richardson 1934, Rigg 1958), the following eight sphagnum bogs were listed: the San Juan peat area, Blakely Island peat area, Killebrew Lake peat area, Orcas N o . 2 peat area, Orcas N o . 1 peat area, C o l d Springs peat area, Constitution N o . 1 peat area, Constitution N o . 2 peat area. Rigg (1925) defines the term sphagnum bog as "that stage i n the physiographic succession o f an area during which its surface is entirely devoid o f ordinary "hard" soil and is composed almost entirely o f living Sphagnum, immediately under which is fibrous brown peat composed mainly or entirely o f partially disintegrated Sphagnum". Understanding Rigg's concept o f Sphagnum bogs was important when trying to interpret his papers or locate some o f his eight bogs. Based on the above information, every attempt was made to visit these unusual bog sites. The results o f these visits are discussed below.  74  The San Juan peat area is about 9.7 k m (six miles) northwest o f Friday Harbor in the Beaverton Valley area on San Juan Island. This deposit was once 6.1 hectares (15 acres) in size, with five acres in Sphagnum.  Rigg (1958) indicates that this bog had been  drained and burned over the years, thus destroying much o f the bog and natural vegetation. When the bog was visited during the present study there was no evidence o f Sphagnum or any o f the typical vascular plants associated with a bog ecosystem, and the area was being used as pasture land. Only one bog was located on Blakely Island, near the o l d Thatcher B a y Post Office. Rigg and Richardson (1934) describe this bog as being the smallest one noted in the islands and, at that time, contained undisturbed native vegetation. In 1958, Rigg again stressed that this bog was still in its natural state. Although three days were spent on Blakely Island, we were unable to locate this bog. A local resident suggested that when the new power line was installed, it may have eliminated the bog. The possible destruction o f this bog is unfortunate because it would have provided an excellent opportunity to study the difference between l o w and high elevation bog systems. The remaining six bogs range from close to sea level to near the top o f M t . Constitution on Orcas Island. The first o f these, Killebrew Lake is located about 1.2 k m (3/4 mile) north o f Grindstone Bay, According to Rigg and Richardson (1934), a bog about 106.6 m by 152.4 m (350 x 500 feet) in size, was present, but it had been burned and pastured, thus destroying most of the natural bog vegetation. In his more recent survey, Rigg (1958) describes the Killebrew Lake area as having 12.1 hectares (30 acres) of peat and about four acres o f Sphagnum bog. During a visit to this lake we did not find the four acres o f Sphagnum bog, but instead we found two very small patches o f Sphagnum henryense mixed in with sedges, and there was no evidence o f vascular plants normally found in bog sites. It is possible that Rigg was mistaken in his 1958 report, or the prior and succeeding damage to the bog has been so severe that the bog has essentially disappeared. 75  Two bogs, (Orcas N o . l , nine acres in size, and Orcas N o . 2, 7.3 hectares (18 acres in size), are located fairly close together in the M t . Woolard area, about 3.2 k m (2 miles) northeast o f the town o f Orcas. A t one time the upper bog (Orcas N o . 2) was mined for peat (pers. comm. M . Grubb), although this ceased when the bog was dammed and made into a lake. This lake now contains a small island covered with Spiraea douglasii (hardhack), Salix sp. (willow), and Pinus contorta (lodgepole pine). A l o n g the margins o f the island, Ledum groenlandicum  (Labrador-tea), Sphagnum fuscum and S.  henryense are found. In the center o f the island, Dicranum scoparium, splendens, and Rhytidiadelphus  Hylocomium  triquetrus were found on the drier sites. The lower bog  (Orcas N o . 1) was destroyed when it was excavated and made into a small pond. O n the southwest slope o f M t . Constitution near the C o l d Springs area, several shallow peat meadows are found. Rigg (1958) refers to this as the Constitution N o . l peat area, and describes it as part Sphagnum bog and part sedge meadow, with some drainage running through it during the winter months. Although Rigg & Richardson (1934) classified these meadows as Sphagnum bogs, they fit the description o f a fen better. Crum (1988) describes a fen as "a grass, sedge or reed-dominated peatland  that  develop under the influence o f mineral-rich, aerated water at or near the surface". A t the present time these meadows are dominated by Carex sp. (sedges) and lack the typical bog type vascular plants. Mosses found in the meadows included Eurhynchium praelongum, Leptodyctium riparium, Fontinalis antipyretica, Sphagnum henryense and S. subsecundum.  The presence o f Sphagnum subsecundum, a species that "grows i n  mineral-rich, open habitats, in sedge mats" (Crum 1988), combined with the other moss species found, further supports the concept that the Constitution N o . l peat area is not a bog, but a fen. Finally, what is likely the only true Sphagnum bog within the San Juan Islands, occurs in Summit Lake near the top o f M t . Constitution. Rigg and Richardson (1934) referred to this location as the Constitution N o . 2 peat area. Near the center o f the lake a 76  floating mat island has been formed and the following mosses were found: Aulacomnium palustre, Bryum uliginosum, Calliergonella  cuspidata, Pleurozium schreberi,  sphagnicola, Polytrichum strictum, Sphagnum capillifolium,  S. fuscum, S.  Pohlia  magellanicum,  S. palustre, S. recurvum, S. rubellum, S. squarrosum, and S. subsecundum. The typical bog vascular plants included: Eriophorum chamissonis and E. gracile (cotton-grass), Drosera rotundifolia (sundew), Vaccinium oxycoccus (wild cranberry), bog billberry, Ledum groenlandicum  (Labrador-tea), Kalmia occidentalis (western swamp laurel) and  several very stunted Pinus contorta (lodgepole pine) trees. This rich and unusual area is located within Moran State Park and is relatively inaccessible. Thus, unless there is a drastic change in the lake water level this area should remain fairly pristine. Two other areas, both on San Juan Island, were found to have Sphagnum growing in them. Dr. Eugene K o z l o f f at the Friday Harbor Biological Laboratories suggested that I examine a small boggy area in the lower portion o f the Beaverton Valley. Rigg (1958) referred to this site as a fresh-water deposit with little or no Sphagnum.  Although  attempts to ditch and drain the area have left fewer than two acres o f Sphagnum, it harbored a rather interesting flora. Mosses collected at this site included Aulacomnium palustre, Calliergonella Rhytidiadelphus  cuspidata, Dicranum scoparium, Polytrichum  triquetrus, Sphagnum capillifolium,  presence o f D. scoparium, P. juniperinum,  juniperinum,  S. henryense and S. recurvum.  The  and R. triquetrus suggests a change in the  hydrology o f the area towards drier conditions. A t the present time this area is under private ownership and is being grazed by cattle. The second Sphagnum location occurs in Sportsman Lake, also located in the Beaverton Valley. When this site was visited a few small clumps o f Sphagnum squarrosum were found growing on floating mats composed o f tree and shrub roots along the west side o f the lake. Rigg (1958) also mentions this site, but again refers to it as a fresh-water deposit with little or no Sphagnum.  77  These bog and bog like habitats and peat deposits are valuable resources for two reasons. First they are the only locations within the islands where the unusual bog type vascular and moss species can survive, and second, these peat deposits hold a wealth o f palynological information. Although the climate is well suited for the development and maintenance o f bogs, they have become a rare resource in Washington State. A l l o f these remaining unusual bog habitats merit further protection.  78  CHAPTER 6 Phytogeography  A complete inventory o f the flora i n an area reveals patterns i n distributions and raises questions as to how these patterns originated. This chapter discusses and interprets the world and the regional Pacific North American distribution patterns for the moss flora in the San Juan Islands. Summaries o f the distribution elements are given in Tables 15. and 16. A complete list of species and their distributions is given in Appendix G and H . Because the islands have been subjected to a great deal o f human disturbance through time, a section o f this chapter focuses on these disturbances and how they have influenced the distribution o f the mosses found in the islands today. The basic goals o f phytogeography are to determine and record species from a fixed location and to map the distribution o f these species. The first task is accomplished through field work and the establishment o f vouchers within a herbarium. Vouchers are an important part of this process as they establish a reliable data base, and make it possible for any errors in determination to be found and corrected. Before any phytogeographical comparisons can be made, floras must be developed. When the floristic information is accumulated from current fieldwork and historical collections, distribution maps can be made, and the species may then be assigned to geographical elements (Stott 1981). Elements are based on recurring patterns. Finally an increasing number o f floristic manuals are needed. According to McLaughlin (1989) " A t the present time the network o f available local floras is inadequate ... More floras need to be compiled - and published - for all floristic areas o f the western United States...". McLaughlin's comments are based on vascular plant floras; extensive bryophyte floras with good habitat data are even more scarce. Several factors influence the distribution o f bryophytes. Historical events, including continental drift, climatic change through time and in space, glaciation or mountain uplift, interacted to determine present bryophyte patterns. Human activities have altered the distribution patterns through transport o f species and the disturbance o f habitat. Ecological factors are summarized by van der Pijl (1972) who states that plant migration can happen only 79  Table 15. The world distributions o f the mosses i n the San Juan Islands No. of Species 43  %of flora 19.19  Circumboreal  92  41.07  Circumtemperate  4  1.78  Circumpolar  29  12.90  North Pacific  4  1.78  Pacific North America - Mediterranean  15  6.69  Western North America -Western Europe  16  7.14  Western North America - Eastern North America  1  .044  Western North America - Eurasia  8  3.57  Widespread  5  2.23  Unclassified  7  3.12  224  100.0  Element Endemic Western North America  Total  Table 16. The Pacific Northwest distributions o f the mosses i n the San Juan Islands. No. of Species 27  %of flora 12.05  47  20.98  Dry Interior - disjunct Coastal  9  4.01  Rocky Mountains - disjunct Coastal  2  .892  Mediterranean Climate  25  11.16  Widespread  114  50.89  Element Coastal Coastal - disjunct Humid Interior  224  Total  80  100.0  when dispersal occurs at the right time and place, and establishment follows on a suitable substratum. Specific habitat and substratum requirements are discussed in Chapter 5. While bryophytes tend to show much wider distribution patterns than do the vascular plants, in general their patterns are similar, suggesting that similar historical and biological events influenced them (Schofield 1992a). The wide bryophyte distribution patterns may have resulted from the supposed ancient age o f the bryophytes, which has provided a long time for them to establish these ranges. Furthermore, the production o f large numbers o f small wind blown diaspores needed for long-range dispersal may be equally important (Schofield 1992a). The dispersibility o f moss spores by wind is well documented, and may occur over short or longrange distances (Crum 1972, Schuster 1983). Establishment and colonization occur mainly in areas that have "unsaturated or unexploited niches" brought about by some catastrophic event, such as volcanism or glaciation (Schuster 1983). Therefore the climatic events o f the Tertiary helped to shape the bryophyte distribution patterns found at present in the world Floristic History According to Axelrod (1958) a Madro-Tertiary Geoflora was present i n southwestern North America by the Early Cenozoic and was located between the Arcto-Tertiary and Neotropical-Tertiary Geofloras. Expansion o f this flora over wide regions began after the Eocene as the drier, warmer climate began to spread. Remnants from this migration can be seen in the vascular flora in the San Juan Islands today, and include the following species: Arbutus menziesii, Berberis nervosa, and Quercus garryana whose fossil equivalents are well represented in the Madro-Tertiary Geoflora. Axelrod (1975) also proposes that some o f the genera may have been derived from the semi-arid regions o f Europe to North America that extended their range via a southerly route that he calls the Madrean-Tethyan link. Examples o f vascular plants that could have utilized this route include: Galium, Juniperus, Pinus, and Quercus. Palynological evidence from the postglacial sediments in the Saanich Inlet, British Columbia also support the early presence o f the Madrean element in this geographic region. L . Heusser (1983) found large amounts o f Quercus pollen in the late Tertiary core samples, indicating that a warmer dry climate had moved into the area. Since this mediterranean type 81  climate developed in the late Tertiary (Axelrod 1973) and the area remained isolated from other areas with mediterranean type climates, Raven (1977) feels that this isolation may be used to explain the high endemism found in California. This same explanation possibly could be used to interpret the large number o f endemic bryophyte species in this climatic zone o f western North America. After the middle Miocene a variety o f species began to enter this region from the north. Wolfe (1969) indicates that even though these species entered from the north they may not have evolved in that immediate area, rather they may have originated in Eurasia and simply migrated through Alaska. Some o f the northern element vascular plant species include Salix hookeriana (Hooker's willow), and Gaultheria shallon (salal), both o f which are common understory species in the San Juan Islands. Alnus sinuata (Sitka alder), a species that is now restricted to a few scattered colonies in the San Juan Islands, and finally Acer circinatum (vine maple) and Rhododendron macrophyllum, both o f which are missing from the island flora (Atkinson and Sharpe 1985), but are widespread on the adjacent mainland. The origin o f the bryophytes o f the northern European element is not clearly understood. These mosses include a number o f widespread wetter climate and mainly forest species such as  Plagiothecium undulatum, Bryum miniatum, Dicranum tauricum and Hookeria lucens (Schofield 1984). A number o f moss species are disjunct between Japan and Pacific Northwest America. A m o n g these are Claopodium crispifolium, Hypnum subimponens, H. dieckii, and Pogonatum contortum. The mosses noted above represent fragments or centers o f three different distribution patterns (Schofield 1965). Floristic Patterns While many floristic distribution patterns have been described in detail for the vascular plants found throughout the world, only a few bryophyte floristic patterns have been described (Herzog (1926), M i l l e r (1982), Schuster (1983), and most recently by Schofield (1992a). Using the bryofloristic kingdoms established by Schofield (1992a) the San Juan Islands are located in Pacific North American Region o f the Holarctic Floristic Kingdom. This kingdom is dominated by a high proportion o f temperate elements, a vast array o f environments, and occupies the 82  largest land mass in the world (Schofield 1992a). Schofield (1992a) attributes the relative uniformity o f the floristic character found in the northern portions o f the kingdom to the recent colonization o f the region following the retreat o f the glaciers, as well as to the wide distribution of similar climates with similar habitats. The southern portion o f this kingdom on the other hand reflects the spatial and climatic isolation that occurred in the region, and led to an increase in endemism. This corresponds nicely with Raven's (1977) views on the high endemism found in California. It is, therefore, not unexpected that 96.4 % o f the San Juan Islands flora reflects this holarctic pattern and that 19.19 % o f the flora is composed o f Western North America endemic species. The Pacific North American Region (see Figure 7.), is one o f the richest bryofloristic regions in North America, with approximately 15 % o f the moss species and 10 % o f the hepatics occurring as endemics. (Schofield 1969, 1980, 1984, 1992a). There are also a number o f genera that reach their greatest diversity in the world, including the following taxa: Buxbaumia (3 spp.), Claopodium (4 spp.), Encalypta (11 spp), Homalothecium  (1 lspp.), and Racomitrium (14 spp.),  (Schofield 1984). Schofield (1992a) divides this region into a northern portion, characterized by a wetter, cooler climate, and a southern portion, with a drier, warmer climate. These two climatic sections, combined with the isolation o f the region by the surrounding high mountains, creates a very distinct bryofloristic region, rich in monotypic genera and abundant, widespread endemic species (Schofield 1992a). Representative monotypic genera include: Alsia, Leucolepis, Pseudobraunia, Plagiomnium  and Rhytidiopsis.  Dendroalsia,  Claopodium bolanderi, Hypnum  circinale,  insigne, and Porotrichum bigelovii are among the widespread endemic species  found in the region. The large number o f distinct and well differentiated endemics suggests that these taxa have been in this region for a long time, and survived the glaciation i n either northern or, more probably, southern refugia.  83  Figure 7. Pacific North American Region, based on Schofield (1992a) and Takhtajan (1986).  The northern portion o f this region contains a large number o f taxa showing disjunctions with Atlantic Europe and south-east Asia, while the southern portion (although not as diverse and less well documented) is characterized by drought tolerant species, and is rich in endemic monotypic genera mixed with disjunct taxa from the Mediterranean region. This southern mediterranean flora may have migrated along the Tethys Sea shoreline, suggesting that it is a relictual fragment in the Pacific Northwest, (Schofield 1992a) or it may have evolved here within the Northwest (Schofield 1994) These drought tolerant species often extend into the northern portion o f this region where suitable habitats exist. The San Juan Islands represent such an area because they harbor a unique blend o f both southern area species (favored by a climate that is a result o f the rainshadow effect created by the Olympic mountains) and the northern area species. Phytogeographic elements were assigned to the 224 species and varieties o f moss found within the San Juan Islands on the basis o f both world distribution and Pacific Northwest distributions, see Tables 17. and 18. These elements were further divided into continuous distribution patterns (including endemics), and disjunct distributions. Godfrey (1977) defines a continuous distribution as "the type o f distribution shown by any taxon occupying more or less all sites suitable for it within the boundaries o f one given region". A n endemic distribution is composed of species that occur in a restricted area usually confined to one region. Disjunct distributions occur when a species occupies two or more areas that are widely separated, yet have suitable habitat occurring in between them (Godfrey 1977). These disjunct patterns have developed as a result o f millions o f years o f floristic change that occurred since the Tertiary period. W o r l d Continuous Distributions Forty one percent o f the species found in the San Juan Islands reflect a Circumboreal distribution pattern. "Circumboreal species are generally considered to have distributions resulting from a modification o f a widespread pre-Pleistocene distribution" (Worley 1972). Representative species include: Aulacomnium  androgynum, Dicranum fuscescens,  antipyretica, and Hylocomium splendens (See Figure 8 ).  85  Fontinalis  Table 17. W o r l d phytogeographic elements i n the Northern Hemisphere. 1. CONTINUOUS DISTRIBUTIONS a. Circumboreal (CB) b. Circumtemperate (CT) c. Circumpolar (CP) d. Endemic Western North America (WNA) e. North Pacific (NP) f. Widespread 2. DISJUNCT DISTRIBUTIONS a. Bipolar (BP) b. Pacific North America - Mediterranean c. Western North America - Eastern North America d. Western North America - Eurasia e. Western North America - Western Europe  Table 18. Pacific Northwest phytogeographic elements. 1. CONTINUOUS DISTRIBUTIONS a. Coastal (C) b. Mediterranean Climate (MC) c. Widespread (W) 2. DISJUNCT DISTRIBUTIONS a. Coastal - Humid Interior (CHI) b. Dry Interior - Coastal (DIC) c. Rocky Mountain - Coastal ( R M C )  86  Figure 8. The Circumboreal pattern in the Northern Hemisphere of the world distribution of Hylocomium splendens. (based on Schofield 1974)  Figure 9. The Circumtemperate pattern in the Northern Hemisphere of the world distribution of Grimmia laevigata, (based on Crum and Anderson 1981, Ignatov and Afonina 1992, Nyholm 1956)  87  Only four species (1.78 %); Didymodon fallax, Grimmia laevigata (see figure 9), G. pulvinata, and Phascum cuspidatum reflect a Circumtemperate distribution pattern. While 29 species (12.9 %) reflect a Circumpolar distribution. A m o n g these species are: Aulacomnium palustre (see figure 10 ), Colliergon giganteum, Leptodictyum Sphagnum squarrosum, and Warnstorfia  riparium,  fluitans.  The North Pacific distribution pattern has been discussed at length by Schofield (1965, 1969, 1980,1984) and appears to reflect three possible patterns o f origin. One theory is that the center o f this distribution was located in Japan and the North American populations reflect remnants o f a wider distribution. Another theory is that hyperoceanic North America was the center o f origin and that fragments extended into Japan. The final theory is that the flora is a fragmented amphi-Pacific distribution. North Pacific element mosses found in the San Juan Islands include the following four species: Claopodium crispifolium, Hypnum dieckii (see figure 11.), H. subimponens, and Pogonatum contortum, and represent only 1.78 % o f the flora. Three of the species, Hypnum dieckii, H. subimponens and Pogonatum contortum are examples of the fragmented amphi-Pacific distribution pattern which "appear to be ancient relict populations o f Tertiary times" (Schofield 1980). Claopodium crispifolium has its greatest distribution in Pacific North America and is disjunct as a few populations in Japan, thus implying an hyperoceanic North American center of origin, and expanded the range along the Aleutian Chain. Widespread or cosmopolitan species are those that occur on all or on most o f the continents throughout the world. The following five species (2.23 %) exhibited this distribution pattern: Bryum argenteum, Ceratodon purpureus, Funaria hygrometirca, purum, and Weissia controversa.  With the exception o f Pseudoscleropodium  Pseudoscleropodium purum all o f the  above species were found on open, inorganic substrata. While Pseudoscleropodium  purum is  considered to be widespread in the Northwest it was found only once on soil mixed in among the grass in an old agricultural field. Pseudoscleropodium  purum (European species) was first reported i n the Pacific  Northwest when Lawton (1960) noted that the species was found in a lawn and flower bed in the Seattle area. A t that time she suggested that the material had been introduced into the area 88  Figure 10. The Circumpolar pattern of the Northern Hemisphere of the world distribution of Aulacomnium palustre. (based on Schofield 1974)  89  by way of imported garden plants. Dickson (1967) announced the discovery o f this species on the islands o f St. Helena and Tristan da Cunha, and suggested that the moss was first introduced to St. Helena from Europe. The introduction to Tristan da Cunha from St. Helena most likely occurred when crates o f young trees packed in P. purum were shipped to the island and the moss was scattered when the trees were planted. It is likely that this moss was introduced to the Shaw Island site by the original property owner, a landscape architect who brought in non-native plant material to landscape around his home. Although this introduction was probably from the Seattle area where P. purum has become fairly well established, and not directly from European material, it has become well accepted that this species has been introduced throughout the world by human horticultural interests and activities. Seven species (3.12 %) were assigned to the unclassified category because either they did not fit into a particular world distribution pattern, or the species concept was not clear; therefore, defining the distribution was not possible. A m o n g the species placed i n this group were Antitrichia  curtipendula, Crumia latifolia, Isothecium myosuroides, Pleuridium acuminatum and  Sphagnum recurvum. For a complete list o f the world distribution patterns see Appendix G .  Endemic Western North American species composed 19.2 % o f the San Juan flora. This phytogeographic element represented the third largest component o f the flora. These endemic species can be divided into three groups based on their distributions. The first group is composed o f those species that are widespread in the islands and elsewhere, such as Anacolia menziesii, Claopodium bolanderi, Eurhynchium oreganum, and Neckera douglasii.  The second  group is composed o f species that are widespread elsewhere, but are uncommon in the islands (probably a result o f not enough suitable habitats), and include Dicranella pacifica, robusta, and Ulota megalospora.  Rhytidiopsis  Finally the third group includes the endemic species that occur  only in the mediterranean climate areas. These drought tolerant species include: Amphidium californicum,  Homalothecium arenarium, H. nuttallii, H. pinnatifidum  throughout the islands), Trachybryum megaptilum, Alsia californica,  (all widespread Dendroalsia abietina (see  Figure 12.), and Fissidens ventricosus (all with limited distributions in the islands.) Schofield 90  (1969) suggests that these endemic species survived during the Pleistocene i n forested areas south o f the glacial boundary, and could have re-entered the region during the hypsithermal interval (Deevey and Flint 1957) when the climatic conditions became warmer.  Disjunct W o r l d Distributions Disjunctions can generally be interpreted as the result o f two possible explanations, longdistance dispersal from a parent population, or persistent fragments from a more continuous historical distribution, that has been disrupted by climatic change, continental drift or a catastrophic event (Godfrey 1977). The bryophyte disjunct patterns used in this paper were recognized and discussed by Schofield and Crum (1972), and Schofield (1969, 1980). A total o f 40 species (17.85 % o f the flora) showed some type o f disjunct distribution pattern, with the largest percentage o f species (13.83 %) disjunct between Western North America and Western Europe or Pacific North America and Mediterranean Europe. The Western North American - Western Europe element is composed o f both oceanic and alpine species that are probably persistent remnants o f an early Tertiary circumboreal flora (Schofield 1969). Representative species include Andreaea megistospora (see Figure 13.), Bryum Hedwigia stellata, and Plagiothecium  undulatum.  miniatum,  Andreaea megistospora occurs from sea level  to subalpine elevations in oceanic regions, and is found in coastal Alaska, British Columbia, Scotland, England, Ireland, Wales, Norway, and in two Washington State locations (Murray 1987). The San Juan collection represents the third location for this species within the state o f Washington, and its occurrence reflects the cool humid environment found on M t . Constitution. Eight species (3.57 %) occur as disjuncts between Western North America and Eurasia. This element represents a mixture o f taxa that were part o f an ancient Temperate amphi-Pacific Tertiary flora and part o f the Circumboreal Tertiary flora (Schofield 1965, 1969, 1984). During the Pleistocene glaciations these taxa probably survived in the unglaciated areas o f Alaska and the Yukon, Coastal Alaska and British Columbia refugial sites or south o f the glacial boundary (Schofield 1980). Migration o f these taxa occurred after the glaciers retreated and the climate became favorable for recolonization. The "persistence o f these species in western North America 91  Figure 12. The Western North American Endemic pattern of the world distribution of Dendroaisia abietina. (based on Manuel 1974 and Schofield 1980)  Figure 13. The Western North American - Western Europe disjunct pattern of the world distribution of Andreaea megistospora. (based on Murray 1987)  92  is related to continued availability o f suitable conditions over a very extended period o f time" (Schofield 1984). Representative species o f this element include Didymodon vinealis var. vinealis, Epipterygium tozeri, Grimmia trichophylla, Orthotrichum hallii (see Figure 14), and Plagiothecium  undulatum.  The recent location o f Orthotrichum hallii in the Altair Mountains o f China by Tan (Lewinsky-Haapasaariana and Tan 1995) is o f particular interest because it was originally known only from western North America. Vitt (1971) considers this species to be a more recently evolved taxon in the North American flora, therefore it probably migrated to A s i a from western North America, although until further populations o f O. hallii are found in China it is difficult to speculate how it got into Asia. The Pacific North American distribution o f O. hallii is discussed later in this chapter. Two species, Hookeria lucens and Pseudoleskea stenophylla follow this disjunct distribution pattern and occur on the mainland fairly frequently, but were not found i n the San Juan Islands flora. The absence o f the Hookeria is not surprising because it is restricted to damp or wet soil in moist wooded areas, and this habitat is not common i n the islands.  Pseuodleskea  stenophylla, on the other hand, is commonly found on the branches and trunks o f living trees, particularly understory species such as Acer circinatum (vine maple) on the mainland. Since vine maple is not found in the islands at the present time it is possible that the Pseudoleskea stenophylla did not extend its range back into the islands for the same reasons that prevented the vine maple from returning. The Western North America - Eastern North America element is represented by one North American endemic species, Heterocladium macounii (see Figure 15 ). Ten other taxa reflect this distribution pattern only in North America but are essentially circumboreal in their world distributions. Schofield (1969, 1972, 1980) discusses this element extensively and proposes that these species were once part o f a widespread flora that became fragmented during the Pleistocene glaciation. It is most likely that these taxa survived south o f the glacial boundaries, migrating northward after the glaciers retreated. Heterocladium  macounii is thought  to be a refugial species in eastern North America because it is found only in unglaciated areas in 93  Figure 14. The Western North American - Eurasia disjunct pattern of the world distribution of Orthotrichum hallii. (based on Lewinsky-Haapasaari and Tan 1995, Vitt 1973)  Figure 15. The Western North American - Eastern North American disjunct pattern of the world distribution of Heterocladium macoimii. (based on Schofield 1985)  94  the southern Appalachians in the east, in the west it is widespread. Schofield (1985a) suggests that the widespread occurrence o f this species could be the result o f the western populations reproducing sexually, giving the species greater heterozygosity and a consequent wider habitat tolerance. Species that have a wider world distribution but show this east-west disjunction in North America include Dryptodon patens, Neckera pennata, Pseudotaxiphyllum papillosa  elegans, Tortula  (all woodland species), and Schistidium maritimum and Ulotaphyllantha  (both  maritime species). While Tortula papillosa is common throughout the northeastern states, Steere (1940) comments that a "single small collection from California, correctly named, has been seen". Schnooberger (1942) also remarks in her paper on the distribution o f Tortula papillosa that the only western locality is in California. Therefore, the four collections o f this species from two o f the San Juan Islands represent the second western North American locality and the first report o f this species for Washington state. Although this epiphytic species is similar to Tortula  latifolia  (a common western species) it can be readily distinguished by its blunt leaves. Neckera pennata presents another interesting distribution pattern, although it does occur in the Rocky Mountains and south into Arizona. The Neckera that is found in that region appears to be different morphologically and in habitat from the eastern material. While this Rocky Mountain type was recognized as N. pennata var. tenera by Flowers (1973) and has been called Neckera oligocarpa by others, at the present time it is usually included with Neckera  pennata.  Further work needs to be done to resolve this taxonomic problem. In the meantime the material collected in the San Juan Islands resembles the eastern material and not this Rocky Mountain type. The bipolar disjunct species were identified in Appendix I only to provide a better understanding o f the overall world distributions for the species found within the San Juan Islands. A representative species reflecting this pattern is Plagiothecium denticulatum  (see  Figure 16). None o f these taxa represent disjuncts from the southern Hemisphere to the San Juan Islands, therefore this bipolar distribution pattern did not influence the moss flora o f the islands.  95  Additional information on bipolar disjunct patterns can be found in the papers by Schofield (1969, 1974, 1980). One o f the more significant elements o f the San Juan Islands moss flora is the Pacific North America - Mediterranean disjuncts. Fifteen species (6.69 %) show this disjunct pattern and all o f them are drought tolerant species that occur within the southern portion o f the Pacific North American Region. This disjunct element has been discussed in detail by Schofield (1969, 1980, 1984, 1988b, 1994), and when combined with the Western North American endemic species found in the islands, it comprises 26 % o f the moss flora. Although the origin o f these disjunct species is not completely clear, two possible scenarios have been suggested by Schofield (1988a, 1994). The first idea is that the center o f origin was located in the Mediterranean region o f Europe and the taxa migrated to Pacific North America. The other scenario is just the opposite, i.e., the taxa originated in the Pacific North American area and migrated to the Mediterranean region. Both o f these ideas suggest a migration route along the coastal shores o f the Tethyan Sea during the late Tertiary. Therefore these disjunct species were probably well established in the Pacific Northwest prior to the last glacial episode, and survived glaciation south o f the glacial boundary in the warmer and drier areas. It is probable that many migrated back into the region during the Hypsithermal Interval. Representative species o f this element include Antitrichia californica (see Figure 17 ), Dicranella  howei, Orthotrichum lyellii, and Scleropodium  cespitans.  Since the San Juan Islands were heavily glaciated during the Pleistocene all o f the vegetation that covers the islands today is composed o f species that have vegetated the islands since deglaciation. The regional floristic patterns reflect those post Pleistocene migrations and they are important in interpreting the moss flora o f the islands at the present time.  96  Figure 16. The Bipolar pattern of the world distribution of Plagiothecium  denticulatum.  (based on Schofield 1974)  Figure 17. The Mediterranean pattern of the world distribution ofAntitrichia (based on Schofield (1980)  97  califomica  Pacific Northwest Distributions  The bryophyte distribution patterns within the Pacific North American Region have been documented and discussed by Schofield (1969, 1976, 1988a). Using a slightly modified version of these geographic distributions the 224 species and varieties o f moss found in the San Juan Islands were assigned to six Pacific Northwest distributional elements. These elements include Coastal (species that occur on the west side o f the Cascade and Coast Mountains), Coastal disjunct to the Humid Interior (species that are found in the Coastal area and then reappear i n the humid areas east o f the Cascade and Coast Mountains), Dry Interior disjunct Coastal (species that occur predominantly east o f the Cascades and Coast Mountains and west of the Rocky Mountains), Rocky Mountain disjunct Coastal (species that have the bulk of their range in the Rocky Mountains but reappear in limited areas in the Coastal area), Mediterranean Climate (includes species that occur in the dry climatic areas west o f the Cascade Mountains), and finally, Widespread (species that occur throughout the Pacific Northwest). Each o f these elements is discussed in detail below. For a complete list o f species and their assigned geographic element, see Appendix H . Table 16. gives a summary o f the number of species found in each element and the percent o f the total flora they represent. A representative species was chosen from each element and a distribution map is provided to show each o f the distributions. Since the Widespread species form the foundation o f most o f the Pacific Northwest flora the large species diversity (114 species or 50.9 % ) , found in the San Juan Island flora was not unexpected, and it is consistent with the moss flora o f British Columbia where just over 100 species belong to this element (Schofield 1976). Species i n this element are usually tolerant o f a wide variety o f environment conditions and are capable o f expanding their range rapidly. Representative species fall into two categories, those that occur on 25 % (or more) o f the islands, e.g. Ceratodon purpureus, Pleurozium schreberi (see Figure 18a.), juniperinum,  P. piliferum and Rhytidiadelphus  Polytrichum  triquetrus, and those that while widespread  elsewhere but occur on only 3.5 % o f the islands. Restricted species include Blindia 98  acuta,  Polytrichum strictum, Sphagnum palustre and S. subsecundum, and reflects the more scarce habitats within these islands. The Coastal element, consisting o f the more humid sites west o f the mountain crest, contains 27 species, or 12.0 % o f the flora. T w o species, Schistidium maritimum and Ulota phyllantha, obligate maritime shoreline species, are i n this element. O n the other hand, Bryum amblydon, a circumpolar species, is usually found in the sub-alpine to alpine areas in this element. On the basis o f where the species may have survived during the Pleistocene glaciation, Schofield (1976) divides this element into the following three components, "species with a more northern distribution, species o f general distribution and subalpine-alpine species". Species of a more northern distribution, such as Andreaea megistospora, may have survived in such refugial sites as the Queen Charlotte Islands, and migrated southward after the glaciers retreated or they could have survived south o f the glacial boundary and subsequently extended northward to sites that are climatically similar to the sites where they survived glaciation. Work by Mathewes (1973) suggests that some o f the species found in this element were present in the southern parts o f British Columbia as early as 10,000 years ago, and these species may have survived i n refugia south o f the glacial boundary. Therefore the species o f general distribution and the subalpine - alpine species most likely returned to the region from southern refugia. Representative species o f the general distribution include Claopodium Dicranoweisia  cirrata, Schistidium maritimum, and Ulota phyllantha.  crispifolium,  (see Figure 18b ). True  subalpine - alpine habitat does not occur within the San Juan Islands, therefore this component is not represented in the islands. Forty-seven species (20.9 % o f the flora) were found within the Coastal - disjunct to the Humid Interior element. The species in this distribution pattern are well represented in the coastal areas, yet reappear in isolated populations on the east side o f the mountains i n areas that are climatically and vegetationally similar to the west side. A substantial number o f these species are Western North American endemics, including the following species: Anacolia 99  menziesii,  Figure 18. The Pacific Northwest distribution of: A . Pleurozium schreberi, a widespread pattern. B . The Coastal distribution pattern o f Ulota phyllantha.  Atrichum selwynii, Claopodium bolanderi (see Figure 19a.), Neckera douglasii, and Rhytidiopsis robusta. Other species, such as Bryum miniatum, Plagiothecium undulatum, and Pogonatum contortum, are disjuntive from western Europe or eastern A s i a . Only nine species, or 4.0 % of the flora represent the Coastal - disjunct to the Dry Interior element. The species o f this element usually occur on mineral soil or on rocks, and survive only in very dry areas; the l o w diversity in this element is not unexpected. Representative species include Desmatodon obtusifolius, Funaria muhlenbergii, Grimmia laevigata, and Trichostomopsis australasiae (see Figure 19b. ). It is reasonable to speculate that these species survived in refugial sites well south o f the glacial boundary and extended northward after the glaciers retreated, possibly as recently as the hypsithermal interval (Deevey and Flint 1957). Atkinson and Sharpe (1985) found, in the vascular plant flora, that "a number o f species that are more common east o f the Cascades have been found i n the San Juans". Representative species include Juniperus scopulorum (Rocky Mountain juniper), Poa scabrella (pine bluegrass), Opuntia fragilis (prickly pear cactus), and Woodsia scopulina (Rocky Mountain woodsia). This demonstrates a disjunct pattern that occurs between the Rocky Mountains and the San Juan Islands. Two species o f moss, Drepanocladus crassicostatus and Orthotrichum hallii seem to be best placed in the Rocky Mountain - disjunct Coastal element. These two species are very limited in distribution and occur on only two o f the islands. Drepanocladus crassicostatus was described by Jannssens (1983a) from living populations and Pleistocene and Holocene age fossil material. He further comments that this new North American taxon is "clearly a western species with arctic-alpine aspect" (Janssens 1983b). The early fossil records and arctic-alpine aspect suggest that D. crassicostatus was part o f the Arcto-Tertiary Geoflora that was widespread at one time. L i v i n g populations o f this species have been recorded from Colorado, Wyoming, British Columbia and the Y u k o n Territory. The recent collection noted above was made during this study and represents the first record for Washington state.  101  Figure 19. The Pacific Northwest distribution of: A. Claopodium bolanderi, a Coastal disjunct to the Humid Interior pattern. B. Trichostomopsis australasiae, a Dry Interior disjunct pattern.  Orthotrichum hallii (see Figure 20a.) as previously noted was once thought to be a Western North American endemic species until it was collected from China by Tan (LewinskyHaapasaari and Tan 1995). Although originally considered to have "evolved relatively recently in situ" (Vitt 1971), it would appear that this species evolved much earlier and possibly was part o f the Arcto-Tertiary Geoflora. This idea is supported by the China collection which places this species into the Western North American - disjunct Eurasia element. The San Juan Islands collection represents the first record for Washington state and suggests an interesting correlation between the disjunct vascular plant species found between the Rocky Mountains and the islands. It is interesting to note that the North American distributions o f O. hallii and D. crassicostatus (living populations only) closely follow the distribution pattern shown by Juniperus scopulorum (see Figure 21 ). The central portion of range for J. scopulorum is located in Colorado, Utah and Wyoming, disjunct populations occur in British Columbia and in the Puget Sound. Most likely the juniper reached its maximum distribution as part o f the Madrean-Tethean Geoflora (Axelrod 1958). Adams (1983) using terpenoid variation i n populations throughout the J. scopulorum range found that "the Manning Pass ( B C ) populations shows closest ties to the Puget Sound populations" and "the greatest differentiation is shown between the Puget Sound populations and the interior populations". Therefore it appears that J. scopulorum originally migrated northward with the Madrean-Tethean Geoflora and was isolated i n the Puget Sound area before the Pliocene. During the Pleistocene glaciation these populations migrated southward (possibly into eastern Oregon) and moved northward during the hypsithermal interval. Since mosses usually move as part o f a whole flora and not as individuals (Crum 1972) it would appear that Orthotrichum hallii, a species exhibiting arid evolutionary morphology (Vitt 1971), may have been well established throughout the region at the same time as the juniper. Therefore it most likely migrated into the San Juan Islands and British Columbia at the same time the junipers did. It must be noted that O. hallii is not an epiphytic species and its relationship to the junipers is purely on the basis o f habitat similarities. A t the present time the disjunct populations o f both the juniper and O. hallii represent the northwestern edge o f their ranges, and while the range o f the 103  Figure 20. The Pacific Northwest distribution of: A . Orthotrichum hallii, a Rocky Mountain disjunct Coastal pattern. B . Alsia californica, a Mediterranean Climate pattern.  Figure 21. The distribution o f Drepanocladus crassicostatus compared to the range o f Juniperus scopulorum. | Drepanocladus crassicostatus (living populations), % Orthotrichum hallii ^ Juniperus scopulorum  and Orthotrichum A  Based on Little Jr. (1971), Vitt (1973) and Janssens (1983a)  105  hallii  (fossil records)  juniper is fairly well established, further fieldwork needs to be done to relate the Rocky Mountain populations with the coastal disjunct ones. It is interesting that the China location o f O. hallii was also "found growing on logs and soil-covered rocks and boulders shaded by a Juniperus grove" (Lewinsky-Haapasaari and Tan (1995) therefore suggesting that there may be a correlation with the juniper migration path in that country as well. Drepanocadus  crassicostatus,  on the other hand, may have extended its range southward  during the cooler climatic periods and was driven northward during the hypsithermal interval. This appears to be the case, as the fossil records point to an arctic-alpine origin for this species. Therefore the southern populations found in Colorado, Wyoming and the San Juan Islands probably represent the southern edge o f its range and are in or near refugial sites. The Mediterranean Climate element contained 25 (11.16 %) species that represent a mixture o f western North American endemic and Pacific North America - disjunct Mediterranean species. This element is found mainly along the coast in western North America extending from southwestern British Columbia to Baja California, and is most abundant in California (Schofield 1988b). Mathewes (1973) reports the presence o f Dendroalsia abietina in lake bottom deposits immediately above the M t . Mazama ash layer, therefore it existed i n the region at about 6.600 yr. B.P.. This strongly suggests that the recolonization o f the region by mediterraean climate type species could have occurred during the hypsithermal interval (Deevey and Flint 1957). Representative species include Alsia californica (See Figure 20b ), Dendroalsia Homalothecium  arenarium, Isothecium cristatum (endemics), Dicranoweisia  abietina, cirrata,  Metaneckera menziesii, Pterogonium gracile, and Tortula laevipila var. meridionalis  (disjuncts).  Species associated with this element reach their greatest numbers on well drained exposed rock outcrop knobs, but they are also well represented as epiphytes and on rock cliffs along stream sides in more open coniferous forests (Schofield 1994). This appears to be the case within the San Juan Islands as over 60 % o f the species occurred in one or more o f the five habitats found in the islands (see Table 19).  106  Table 19. The relationship between the number o f mediterraean climate species found in climate found i n each habitat. Total N o . o f Species 16  % o f Total N o . o f Species  Meadows & Ridges  15  60.0  Wetland  15  60.0  Outcrops  16  64.0  Disturbed Sites  10  40.0  Woodlands  19  76.0  Habitat Maritime  107  64.0  One species found in this element merits extra discussion. Tortula laevipila var. meridionalis (a disjunct species) was found on several o f the islands and its collection represents a new record for the United States. Although there has been significant discussion in Europe over the validity o f this variety, Anderson et al. (1990) have accepted it as a valid North American taxon. This species is epiphytic on Acer macrophyllum  (bigleaf maple) and Quercus  garryana  (Garry oak) and two recent collections made on oak trees in the Tacoma, Washington and the Willamette Valley, Oregon areas strongly suggest that this species w i l l ultimately be found throughout the entire Garry oak distribution. This pattern has already been demonstrated with Dendroalsia  abietina (an epiphyte on both maple and oak) when the two distributions are  compared (see Figure 22 ). Only additional field work throughout the region w i l l clarify understanding o f the distribution o f Tortula laevipila var.  meridionalis.  Although a number o f different phytogeographical elements are represented within the islands, three points must be considered when interpreting these patterns. The first o f these is that the entire flora o f the islands represents the result of post Pleistocene migration and the islands presented no refugial sites during the glacial period. Second, when making comparisons between the moss flora o f the San Juan Islands with the immediate adjacent areas it must be understood that a sound data base for most o f the mainland and adjacent islands is lacking. Finally, the environmental destruction that has been ongoing from the time humans entered the region, continues to influence the number of species that occur in the area. These points are discussed in the following sections o f this chapter. Post glacial migration into the islands could have begun as early as 13,500 yr. B . P . (Deither et al. 1995). when the higher areas in the islands were exposed above the ice. These open treeless areas would have provided suitable habitat for moss spores or fragments to travel, settle and become established. Waitt et al. (1983) suggests that the alpine glaciers in the region were greatly diminished around 17000 -18000 yr. B.P. therefore it is possible that vegetation on these exposed higher areas on the mainland could have provided a source o f moss spores and fragments to enter the islands. W i n d dispersal of spores has been well established by Bergeron 108  Figure 22. The Distribution of Dendroalsia abietina Quercus garryana. ^  compared to the range of  (1944), Crum (1972), Schofield and Crum (1972) and more recently by van Zanten et al. (1981). M i l l e r et al. (1976) demonstrated that "some gametophyte fragments are able to function as propagules in nature and that wind dispersal and establishment o f bryophytes by vegetative means may be routine in arctic latitudes". Since the climate in the region could have generated wind patterns that could have distributed spores and fragments from the mainland into the islands it is suggested that some o f the more northern element species may have arrived i n the islands sometime after 13,500 yr. B . P . The continued existence o f these species, especially on the summit o f M t . Constitution is the result o f the cooler, wetter climate that exists there now. Since "...mosses have moved in the past as members o f whole floras i n migration, not as individuals and not aimlessly, but along natural migration routes..." (Crum 1972) it is safe to assume that the remainder o f the flora migrated into the islands most likely from a southern refugial site, during the hypsithermal interval. Lawton (1971) defines the Pacific Northwest as including Washington, Oregon, Idaho, western Montana, Wyoming and north to about the 52nd parallel, with a total moss flora o f "598 species and varieties, attributed to 44 families and 156 genera". In order to present a comparison o f the San Juan Island flora with the Pacific Northwest region defined by Lawton (1971) Table was developed. Checklists for each of the above states and province were available except for Washington considered as a separate unit, therefore Washington as an independent flora is not included in Table 20. Considering the small area of the islands (172 k m ), they have a very diverse moss flora, consisting o f 224 species and varieties. The islands possess 37. 4% o f the species found throughout the Pacific Northwest and 20.3% o f the North American flora. The l o w diversity o f species found in relation to the British Columbia flora is the result o f the greater habitat diversity that occurs throughout the province as well as its larger area. O n a more local level the G u l f Islands o f British Columbia, (located just north o f the San Juan Islands i n the southern portion o f the Strait o f Georgia), contain 114 species and varieties although additional field work w i l l probably add species to the list. It is reasonably safe to assume that the two floras would be 110  Table 20. San Juan moss diversity comparisons to other areas. Location North America  N o . o f Species 1103  % o f Flora 20.3  British Columbia  714  31.4  Pacific Northwest  598  37.4  California  495  45.2  Oregon  441  50.7  Montana  408  54.9  Wyoming  315  71.1  Idaho  257  87.1  San Juan Islands  224  100.0  Anderson et al. (1990), Schofield (1992b), Lawton (1971), Norris (unpublished), Christy et al. (1982), Elliott (unpublished), Eckel (1996), McCleary et al. (1971).  Ill  almost identical except for the Sphagnum bog species which do not occur i n the G u l f Islands. In general the moss flora o f the San Juan Islands is diverse but it is always possible to add additional species to a list when more field work is done. In order to assess o f the frequency for the species within the islands the data base was sorted to provide the total number o f records and number o f islands on which each species was found. Frequency was determined by dividing the number o f islands where a species occurred by the total number of islands visited. The results are given in Appendix I . A species was considered to be common i f it occurred on 50% or more o f the islands, and six species were found on 90 % or more o f the islands. Representative species in this group include Dicranum scoparium, Polytrichum junipehnum,  and Eurhynchium oreganum, all o f which are consistently  widespread on the west side o f the Cascade and Coast mountains, therefore it was not unexpected that they would appear as very common. Rare or infrequent species included any species that occurred on five or fewer o f the islands. Species in this category include Crumia Fissidens grandifrons and Pseudoscleropodium  latifolia,  purum, and the Sphagnum species. It is  important to caution that the "degree o f rarity" reflected by some species in the islands may not be rare on the mainland or adjacent Vancouver Island. This is demonstrated with P. purum and the Sphagnum% which are restricted in the San Juan Islands but are more prevalent on the mainland. Crumia latifolia and Fissidens grandifrons (both obligate calcicoles) are restricted to a few sites in the islands and on the mainland. Therefore a knowledge o f the floras throughout the region is important before a species can be listed as rare on a state or regional list. Although the role o f disturbance is discussed in previous chapters it can be segregated into human and non-human impacts, both play a role in determining where a species may occur. Human impacts are generally apparent and include agriculture, building, recreation, and timber harvest etc. Housing and the development that is associated with it poses the greatest current threat to most habitats on the islands. It is o f significance that a number o f property owners are becoming aware o f the need to preserve their land and are acting to prevent future destruction by setting up conservation easements, etc.. Several organizations have been established i n the 112  islands to assist land owners with this process. Areas already under protection such as the United States Fish and Wildlife Refuges and The Nature Conservancy are certainly beneficial, provide habitat where public access is limited, and offer the best areas for survival for a number o f the species found within the islands. Finally the State Parks and National Park while allowing public access have taken a greater interest in protecting from excess use any critical environments placed in their care. Probably one o f the best examples o f human impact on the distribution o f a species is demonstrated by Dendroalsia abietina, an epiphytic species usually found on Acer  macrophyllum  (bigleaf maple) or Quercus garryana (Garry Oak), and sometimes on rock. It is possible that the volume o f trees cut down over the years to fuel the lime kilns, "feed" the steamers that moved about the islands and the clearing o f land for agricultural purposes may have substantially reduced the available habitat for this species. A t the present time Dendroalsia is found only on very old oak trees scattered on four islands but it may have been more widespread before people become well established in the islands and removed old oak trees during this settlement. Finally logging in the islands occurred extensively on Orcas and Waldron Islands; true old growth forest ecosystems are now absent in the islands. The loss o f this type o f habitat may explain why such dense woodland species as Buxbaumia piperi and Tetraphis geniculata were not found in the islands. The role o f non-human disturbance on the moss flora may be explained through in two scenarios. The first o f these is related to the impact o f birds nesting on some o f the smaller islands. Goose Island demonstrates this well, with a flora o f only 10 species. Human impact on this island is limited related to its inaccessibility and ownership, but it is a major nesting site for gulls. Therefore it is assumed that the concentration o f bird droppings prevents the establishment of a large number o f moss species or severely limits their distribution on the island.  Schistidium  maritimum, a common species o f rock in the maritime sub-habitat throughout the islands was found in only one spot covering an area about 2 cm. in diameter. Additional field work on other islands used as nesting sites needs to be completed to further support this assumption, but at the 113  present time it appears to be a reasonable explanation for the low number o f species found on Goose Island. Another non-human form o f disturbance that occurs when a north-easterly storm brings very strong winds into the islands. These storms often result in a tremendous amount o f blown down timber, this changes the light, temperature and humidity within the forest ecosystems affected. When these changes affect the coniferous woodlands, the mosses such as undulatum and Rhytidiadelphus  Plagiothecium  loreus usually cannot survive. This phenomenon was observed  on Orcas Island in Moran State Park and on Jones Island where a recent north-easterly storm damaged a large volume o f timber. In conclusion, The bulk o f the San Juan Islands moss flora is composed o f circumboreal species that reflect a once more widespread Arcto-Tertiary flora. During the Pleistocene these islands were completely denuded several times. The flora and regional Pacific Northwest phytogeographical patterns that exist now are composed o f species that have migrated back into the region, including the islands. This recolonization could have begun as early as 13,500 yr. B . P . (Deither et al. 1995) but the bulk o f the flora probably returned during the hypsithermal interval described by Deevey and Flint (1957). During the Pleistocene glaciation, areas south o f the glacial boundary possibly served as refugial sites, although some species might have survived in northern refugia. A decrease i n the amount o f Quercus pollen in the Saanich Inlet (British Columbia) deposits occurred about 2000 years ago. A t the same time an increase i n the ThujaChamaecyparis  pollen levels was noted (L. Heusser 1983). This implies that the climate in the  region was changing from warm and dry into a cooler, wetter period. During this cooling trend the southern element appears to have retreated, with fragments remaining only in those areas where conditions were favorable. Thus the San Juan and adjacent islands represent a "modern" refugium for southern, mediterranean type climate species. It is hoped that the people who inhabit this unusual ecosystem w i l l continue to become aware o f its uniqueness and w i l l support its preservation. 114  CHAPTER 7 Taxonomic Considerations  The known moss flora o f the San Juan Islands is composed o f 224 species and varieties representing 33 families and 97 genera. Complete lists o f these species and varieties are provided in Appendix J and K . In addition to the 224 species and varieties collected during the present study, four historical taxa were found in various herbaria throughout the region. Identification keys were prepared to genera and species, and vouchers have been deposited in the University o f British Columbia and University o f Washington herbaria. A list o f taxa not found during this study, but suspected to occur i n the islands is provided i n Appendix M . These taxa were chosen because they are found on the adjacent islands or are common i n similar habitats on the mainland. Distribution maps are provided for all species collected during this study i n Appendix N .  Preparation o f Keys  The dichotomous keys to genera and species were developed from personal observations made during the process o f identification. In addition the following authors were consulted i n order to clarify difficult taxa. Buck and Norris(1996) C r u m (1984) C r u m and Anderson (1981) Crundwell and N y h o l m (1977) Flowers (1973) Frisvoll(1982, 1988) Horton(1982, 1983) Hoisngton(1979) Ireland (1969, 1970, 1971,1982) Ireland and Spence (1987) Janssens (1983a, 1983b) Koponen(1974)  Lawton (1965, 1971) Murray (1987) Noguchi (1987-1994) Robinson (1970) Schofield (1969, 1970) Sharp etal. (1994) Shaw (1982) Spence(1988) Vitt (1973) Vitt and Buck (1992) Welch (1960) Zander(1978)  These keys are designed to be used in conjunction with Lawton's Moss Flora of the Pacific Northwest (1973) and in most cases where measurements are provided i n the keys they 115  have been taken from her book. Because the species listed i n the keys reflect the nomenclature o f the current North American checklist (Anderson et al. 1990), a list o f synonyms is provided for species that are different or missing from Lawton's text (Appendix L . ) . Whenever possible, the keys are based on gametophytic structures. Sporophytic characters are used when it is difficult to determine the species on gametophytic characters alone. Whenever ecological information is useful, it is provided. Use o f these keys requires the use o f both dissecting and compound microscopes and it is important to have reliable ecological information available for each collection, because it is often needed in order to make a determination. Problematic Taxa Atrichum P. Beauv. Both Atrichum selwynii and A. undulatum were collected from the islands with A. selwynii the more common species found. Distinguishing between these two species is strongly based on sexuality, therefore young material can be difficult to distinguish. Ireland (1969, 1971) separates sterile material on the basis of leaf shape and the position o f the dorsal teeth on the surface o f the lamina. In A. selwynii these teeth are distinct and in several diagonal rows. A. undulatum, on the other hand, had teeth that are few or lacking and do not form distinct rows. Unfortunately sexuality can often be a problem, while A. selwynii is always dioicous, A. undulatum is usually monoicous, but rarely dioicous, therefore the gametophytic characteristics become more important in identification.  Dicranella  howei Ren. & Card.  Dicranella  howei is distinguished from D. varia by Crundwell and N y h o l m (1977) based  on leaf and exothecium characters.  Brachythecium  asperrimum (Mitt.) Sull.  This species is considered as distinct from B.frigidum  by Hoisington (1979) and can be  distinguished from B. frigidum on the following characteristics. B. asperrimum has a less 116  defined alar cell region, the leaf tips are usually twisted and the species is always found in well drained habitats and forms loose open mats, it is commonly epiphytic or on logs. B. frigidum has a well defined alar region, often enlarged cells extending across the base o f the leaf, is usually found in wet habitats, often submerged, and never forms loose open mats; it is commonly terrestrial on raw mineral soil. Dicranum howellii Ren. & Card. Lawton (1971) distinguishes D. howellii from D. scoparium on the basis o f the inner perchaetial bracts being gradually acuminate, while they are abruptly acuminate in D. scoparium.  Although this taxon is treated by Anderson et al. (1990) as a valid species, careful  comparison o f the perichaetial bracts in both species demonstrates that D. howellii should be treated within D. scoparium.  It is therefore treated as D. scoparium here.  Fissidens bryoides Hedw. complex Lawton (1971) combined Fissidens limbatus with F. bryoides, therefore placing all o f the material through the Pacific Northwest under F. bryoides. Flowers (1973) recognizes F. limbatus as a distinct species but comments "The differences usually cited in Fissidens  bryoides,  F. limbatus, F. sublimbatus are so slight and intergrade so freely that it is difficult to admit them as full species". Pursell (who is working on Fissidens for the Flora North America project), after examining my material has assigned the following three names to the specimens; F. bryoides var. bryoides, F. bryoides var. viridulus and has placed F. limbatus into F. crispus. The two varieties o f Fissidens bryoides have been included in this study, but the name F. limbatus has not been altered to F. crispus, since the substantiating data are unpublished.  Hedwigia stellata Hedenas Recent studies by Buck and Norris (1996) have concluded that there are several species of Hedwigia in North America. Upon Buck's suggestion, reexamination o f all o f the San Juan material called H. ciliata was found to be H. stellata. H. stellata is distinguished from H. ciliata based on the number o f papillae located on the upper laminal cells. H. ciliata has 2-4 papillae 117  cells (rarely 1) on the upper laminal cells, while H. stellata has 1 or 2 papillae on the upper laminal cells.  Racomitrium heterostichum  (Hedw.) Brid.  Group  The Section Laevifolia (heterostichum) group is an extremely variable group o f taxa, that are difficult to discriminate. Frisvoll (1988) revised this group using the following taxonomic characters that he felt were stable; alar cells and supra-alar marginal cells, costa structure, leaf margin and perichaetial leaf structure. Although he comments in his treatment that "It has proved difficult to make reliable keys for the taxa in the section, primarily because they are all easily modified by differences in environmental factors" (Frisvoll 1988). Using his treatment, three species were recognized, R. lawtonae, R. paciflcum and R.  occidentale.  R. pacificum has ovate-lanceolate leaves, unistratose margins, and lacks a hairpoint, the upper laminal cells are smooth and isodiametric. It could be confused with R. aquaticum and R. varium but both differ by having multipapillose leaf cells (Ireland and Spence 1987, Frisvoll 1988). R. lawtonae has long lanceolate leaves with unistratose margins, the costa is deeply channeled and ends in a long clear hairpoint that is very decurrent extending down the margins o f the leaf as a hyaline area. Leaf cells just below the apex are elongate, rather than isodiametric or mixed short and long (Ireland 1970, Frisvoll 1988).  R. occidentale has ovate-lanceolate leaves with bistratose margins, the hairpoint is either short or long. The most significant feature o f this species is the bulging cells on the dorsal side of the costa (Frisvoll 1988). This species cannot be confused with Dryptodon patens which has well defined "wings" on the dorsal side o f the costa. Collections that did not fit into any o f the above species were placed into Racomitrium hetrostichum sensu lato. The margins o f the lamina were usually unistratose or occasionally had bistratose spots. The hairpoint often was decurrent but never formed the long narrow bands  118  down the margin o f the leaf, as in R. lawtonae. This group o f highly variable o f individuals may contain some material o f true R. heterostichum but it was very difficult to sort these out.  Historical Taxa Although search o f herbarium records in the region resulted i n a list o f 45 species that represented the only available record for an island (see Appendix A . ) . Four o f these species, Distichium capillaceum, Hygrohypnum bestii, Roellia roellii and Tayloria serrata were not relocated on any o f the islands, therefore they represent the only records for these species. Although with the exception o f Roellia roellii (a species usually found only at higher elevations on the mainland) these species may be revealed during future fieldwork, they have been excluded from the key until new populations can be located.  Taxa N e w To Washington State During the present study, four taxa new to Washington State were found, one represents a new record for the United States and the second record for North America. A careful search throughout regional herbaria and the literature confirmed that these were obviously new records. Careful comparison to known vouchers assisted in verifying their identity and the Drepanocladus  crassicostatus was determined by Janssens who originally described the species.  The phytogeographic implications of these species are discussed in Chapter 6.  Drepanocladus  crassicostatus  Janssens  This species is distinguished by the very thick and wide costa at the base o f the main stem leaves, and the leaf margins are lightly to sharply denticulate (Janssens 1983a).  Orthotrichum hallii Sull. and Lesq. in Sull.  This species is found on calcareous rock and has leaves that are bistratose in the upper half, sometimes with unistratose streaks. The sporophyte is immersed to 1/2 emergent and has immersed stomata. It can be distinguished from O. anomalum, which has an exserted 119  sporophyte, and leaves that are unistratose. It is readily separated from O. rupestre by the superficial stomata found in the sporophyte. (Vitt 1973)  Tortula laevipila var. meridionalis (Schimp.) Wijk and Marg. This species is identified on the basis o f the costate leaf-like gemmae that are clustered at the top o f the stem. It can be separated from T. laevipila var. laevipila by the gemmae and the costa which is excurrent as a short hairpoint or is absent (Barkman 1963, Smith 1978).  Tortula papillosa W i l s . ex Spruce This species has broadly obovate-spathulate leaves, with an excurrent costa that forms a smooth hairpoint. Multicellular, spherical or globose propagula are borne on the upper surface of the leaf. This species is easily distinguished from T. latifolia which has an obtuse apex, and lacks the hairpoint (Smith 1978, Crum and Anderson 1981).  120  K E Y TO GENERA 1. Branches clustered i n fascicles; leaf cells composed o f a network o f large hyaline cells and smaller green cells; found in bogs, fens, and along lake margins Sphagnum 1. Branches not clustered i n fascicles; leaf cells never having large hyaline or green cells; found on various substrata 2. 2. Leaves complanate  3.  2. Leaves not complanate, arranged in more than two rows  7.  3. Leaves undulate; stems with or without paraphyllia; plants never aquatic  4.  3. Leaves not undulate; stems without paraphyllia; plants usually aquatic or i n damp areas  6.  4. Stems with abundant paraphyllia; leaf costa strong, to the middle  Metaneckera  4. Stems without paraphyllia; costa lacking or short  5.  5. A l a r cells differentiated, forming several rows o f decurrent rectangular cells ... Plagiothecium  undulatum 5. A l a r cells not differentiated, irregular, quadrate:  6.  6. Leaves with a vaginate lamina; median cells rounded-hexagonal; never undulate  Fissidens 6. Leaves never with a vaginate lamina; median cells elongate; undulate  Neckera  7. Plants acrocarpous, stems and branches erect; sporophytes develop terminally on the main stem 7. Plants pleurocarpous, stems branched, creeping; sporophytes develop on the lateral " ~ 8. Leaf cells smooth  8.  80. 9.  8. Leaf cells papillose  44.  9. Plants dendroid  10.  9. Plants never dendroid  11 •  10. Paraphyllia abundant on stems  Climacium  10. Paraphyllia never found on the stems  Leucolepis 121  11. Leaves with lamellae on the upper surface  12.  11. Leaves without lamellae  15.  12. Lamellae less than 20; leaves undulate, margins bordered  Atrichum  12. Lamellae greater than 20; leaves never undulate, margins not bordered 13. A p i c a l cells o f lamellae smooth, sporophyte angular, never cylindrical  13. Polytrichum  13. Apical cells o f lamellae papillose or smooth, sporophyte cylindrical, rarely angular  14.  14. A p i c a l cells o f lamellae not, papillose, i f smooth then leaves are strongly contorted when dry Pogonatum 14. A p i c a l cells o f lamellae pyriform, papillose, never smooth  Polytrichastrum alpinum  15. Peristome composed o f 4 teeth; disk-shaped, multicellular stalked gemmae i n a cupshaped apical rosette, on rotten logs Tetraphis pellucida 15. Peristome teeth more than 4, or absent  16.  16. Leaves linear-setaceous; sporophyte elongate-pyriform and nodding, with a long slender neck Leptobryum pyriforme 16. Leaves not linear-setaceous  17.  17. Median leaf cells oblong-hexagonal or linear  18.  17. Upper leaf cells isodiametric, rounded quadrate to quadrate ( i f elongate then the leaves are matted with rhizoids)  27.  18. Leaf margins incurved  19.  18. Leaf margins plane, not incurved  23.  19. A l a r cells inflated, forming distinct reddish-brown groups; sporophytes exserted  20.  19. A l a r cells not differentiated or inflated; sporophyte immersed or exserted, teeth present or absent, usually on dry soil  21.  20. Basal cells above inflated cells short, almost quadrate; usually on wet rock.. Blindia acuta 20. Basal cells above inflated cells rectangular to linear; never on wet rock 21. Sporophyte cleistocarpus, immersed on a very short seta 122  Dicranum  Pleuridium acuminatum  21. Sporophyte cleistocarpus, exserted  22.  22. Leaves erect, narrowly lanceolate, usually ending i n a narrow awn; peristome teeth finely divided Ditrichum 22. Leaves lanceolate-subulate, variable, spreading or squarrose; peristome teeth broad  Dicranella 21>. Leaves narrowly lanceolate or lanceolate-subulate  24.  23. Leaves broad, never lanceolate-subulate  25.  24. Leaves erect, narrowly lanceolate, ending in a slender apex, peristome teeth narrow and finely divided Ditrichum 24. Leaves spreading or squarrose, lanceolate-subulate, peristome teeth broad or absent  Dicranella 25. Leaf margin never bordered, median leaf cells long and linear; costa never excurrent  26  25. Leaf margin bordered or not; median leaf cells rhomboidal to hexagonal; costa + excurrent 27. 26. Upper median cells long and linear  Pohlia  26. Upper median cells rhomboidal to hexagonal-oblong  Bryum  27. Leaves bordered with reddish linear cells; costa to just beyond the middle  Epipterygium  tozeri 27. Leaves + bordered, never with reddish linear cells; costa strong, percurrent to excurrent  Bryum 28. Leaves broad, oblong-lanceolate, oblong-elliptic to ovate-lanceolate  29.  28. Leaves linear-lanceolate, never broad, oblong-lanceolate or elliptic  33.  29. Leaf margins bordered and toothed with single or double teeth  30.  29. Leaf margin + bordered, never toothed  31.  30. Leaf border unistratose; teeth single and to the base o f the leaf; on soil, rotten logs or trees  Plagiomnium  30. Leaf border multistratose, teeth double and not extending to the base o f the leaf, usually only on soil or soil over rock ....Mnium  123  31. Leaves oblong-lanceolate, margins not bordered, slightly contorted when dry, on soil  32.  31. Leaves broad, obovate + orbicular, margins with a multistratose border, often strongly contorted when dry, on rotten logs or sometimes soil Rhizomnium 32. Leaf margin entire, sporophyte immersed; peristome teeth absent  Physcomitrium pyriforme 32. Leaf margins entire in the lower half, + serrulate i n the upper half; sporophyte exserted, peristome teeth present Funaria 33. Median leaf cells smooth or slightly papillose  34.  33. Upper cells smooth, never papillose  37.  34. Median leaf cell walls sinuous; leaf apex + a hairpoint  Racomitrium  34. Median leaf cell walls never sinuous; leaf apex without a hairpoint 35. Leaves matted with reddish-brown rhizoids; sporophyte exserted  35. Anacolia menziesii  35. Leaves not matted with rhizoids, sporophyte exserted or immersed  36.  36. Leaves linear-lanceolate, acute, entire or serrulate at apex; sporophyte exserted  Cynodontium jenneri 36. Leaves lanceolate to ovate-lanceolate, usually with a hairpoint; sporophyte immersed  Schistidium 37. Leaves abruptly narrowed from an obovate base; sporophyte very strumose  Oncophorus  wahlenbergii 37. Leaves never abruptly narrowed from an obovate base; sporophyte not strumose  38.  38. Plants on soil, usually in disturbed sites; leaves slightly twisted or contorted when dry, often reddish-purple; sporophyte strongly ribbed when dry Ceratodon purpureus 38. Plants on rock or trees, not i n disturbed sites; leaves strongly to slightly contorted when.... dry, green to blackish-green  39.  39. Plants usually with gemmae; leaves strongly twisted and contorted when dry  40.  39. Plants without gemmae; leaves not strongly twisted and contorted when dry; always on rock  41.  40. Lower leaf cell walls thick, slightly sinuous; always on rock cliffs  Grimmia  torquata 124  40. Lower leaf cell walls not thick, never sinuous; usually on coniferous trees or occasionally on rock Dicranoweisia cirrata 41. Leaves with prominent dorsal wings on the outer surface o f the costa  Dryptodon patens  41. Leaves without prominent dorsal wings on the costa 42. Leaf apex coarsely toothed; calyptra mitrate  42. Ptychomitrium gardneri  42. Leaf apex never coarsely toothed; calyptra not mitrate  43.  43. Calyptra large, campanulate and persistent; lower leaf cell walls lightly sinuous and thickwalled Coscinodon calyptratus 43. Calyptra smaller, not persistent; lower leaf cells sometimes sinuous and ± thickwalled  Grimmia  44. Leaf cells weakly papillose, usually appearing smooth  45.  44. Leaf cells strongly papillose with superficial papillae or by projecting cell ends  46.  45. Leaves lanceolate-subulate, usually ending in a slender apex; sporophyte exserted, peristome teeth finely divided Ditrichum 45. Leaves lanceolate, never subulate; sporophyte immersed, cleistocarpous  Pleuridium  acuminatum 46. Upper leaf cells papillose by projecting cell ends  47.  46. Upper leaf cells papillose with one or more papillae per cell, or cells mammillose  49.  47. A l a r cells inflated, often orange or brown; leaves sometimes with several dorsal ridges; on a wide variety o f substrata  Dicranum  47. A l a r cells not inflated or differentiated; leaves without several dorsal ridges 48. Upper cells linear-oblong, margins recurved (occasionally plane), serrate  48. Philonotis  48. Upper cells not linear-oblong, margins plane or incurved; papillose on a subulate awn  Trichodon cylindricus 49. Propagula usually present  50.  49. Propagula, gemmae or brood bodies absent  57. 125  50. Propagula fusiform, on the tip o f a pseudopodum, very common  Aulacomnium  50. Propagula filamentous, multicellular and leaf shaped, or spherical 51. Propagula  51.  filamentous  52.  51. Propagula multicellular, globose or leaf shaped  55.  52. Filamentous propagula, branched and composed o f papillose cells  Encalypta procera  52. Brown multicellular gemmae usually very common  53.  53. Leaf apex covered with reddish-brown gemmae, or apex forming a rough knob i f gemmae have been broken off; on rocks or trees along the coast Ulota phyllanta 53. Abundant brown multicellular gemmae attached to the surface o f the leaf  54.  54. Leaves strongly twisted and contorted when dry; peristome lacking; on trees or rocks  Zygodon viridisimus var. rupestris 54. Leaves not contorted when dry, peristome always present 55. Propagula leaf-shaped, in clusters in the leaf axils  Orthotrichum  Totrula laevipila v. meridionalis  55. Propagula multicellular, globose or spherical, on the upper surface o f the leaf or in the leaf axils  56.  56. Propagula globose or spherical, on the upper surface o f the leaf; always found on trees  Tortula 56. Multicellular propagula on branched stalk-like structures in the axils o f the leaves; always found on wet rock Dichodontium pellucidum 57. Plants always found on wet calcareous rocks or soil  58.  57. Plants not growing in wet calcareous sites  61.  58. Leaves oblong-spatulate, bordered by 4-7 rows o f large orange cells  Crumia latifolia  58. Leaves linear-lanceolate, not bordered  59.  59. Leaves with sharp marginal teeth, often recurved at the base; peristome teeth present  Eucladium verticillatum 59. Leaves without sharp marginal teeth at the base; peristome teeth absent 60. Leaf margin plane, not recurved  60.  Gymnostomum aeruginosum 126  60. Leaf margin recurved on one or both sides; operculum commonly attached to the columella  Hymenostylium recurvirostre  61. Leaf margin involute, never revolute  62.  61. Leaf margins plane or revolute  66.  62. Plants green to dark green; sporophyte cylindric, + peristome teeth  63.  62. Plants reddish black to dark brown; very small, always on siliceous rock; sporophyte ellipsoid with 4 longitudinal slits, peristome lacking Andreaea 63. Costa glossy, excurrent to a short point; leaves strongly crisped when dry  Weissia controversa 63. Costa not glossy, percurrent to excurrent, apex apiculate or piliferous; leaves moderately to strongly crisped when dry 64. 64. Calyptra large and persistent, covering half or all o f the sporophyte; leaves ligulatespatulate, with numerous papillae on the upper surface; moderately crisped when dry  Encalypta 64. Calyptra smaller, not persistent; leaves linear-lanceolate, median cells mammillose or with several small papillae; leaves strongly twisted and crisped when dry 65. 65. Median leaf cells strongly mammillose; costa broad, ending in or near the apex  Timmiella crassinervis 65. Median leaf cells with several small papillae per cell; costa strong, percurrent or excurrent; basal cells hyaline, extending " V " shaped up the margins Tortella 66. Leaf margin or apex serrate or toothed  67.  66. Leaf margin or apex not serrate or toothed  70.  67. Upper cells mammillose  68.  67. Upper cells papillose, not mammillose  69.  68. Leaves ovate-lanceolate, apex finely or coarsely serrulate-denticulate, on wet soil or rocks  Dichodontium pellucidum 68. Leaves linear-lanceolate from an oblong, orange-brown expanded hyaline base; upper margins coarsely toothed with sharp teeth; on soil Timmia austriaca  127  69. Plants always on siliceous rock; sporophyte ellipsoid with 4 slits, peristome lacking; reddish black and very small  Andreaea  69. Plants on rock, rock cliff faces or on trees  70.  70. Plants usually on rock, rock cliff faces or on trees  71.  70. Plants usually on soil, sometimes on rock or trees  74.  71. Plants on rock cliff faces, never found on trees; strongly crisped and contorted when dry 72. 71. Plants on rocks or trees; + crisped and contorted when dry  73.  72. Plants matted with papillose reddish-brown rhizoids; sporophyte globose, zygomorphic exserted, peristome teeth present Bartramia pomiformis 72. Plants not matted with rhizoids; sporophyte immersed, peristome teeth lacking  Amphidium 73. Plants strongly crisped and contorted when dry; sporophyte stomata superficial  Ulota  73. Plants never strongly crisped and contorted when dry; sporophyte stomata superficial or immersed  Orthotrichum  74. Leaf median cells densely papillose with C - or O-shaped papillae  75.  74. Leaf median cells papillose with several papillae, never C - or O-shaped  78.  75. Sporophyte immersed to emergent, peristome teeth absent; always on soil  Phaseum cuspidatum 75. Sporophyte exserted, peristome teeth present; on soil or trees  76.  76. Leaves oblong to ligulate-spatulate, basal cells hyaline, smooth, extending almost to the middle or beyond; on trees and soil Tortula 76. Leaves linear-lanceolate, oblong-ligulate, never spatulate, basal cells elongate, clear but not extending up the margins o f the leaf; on soil 77. 77. Leaves linear-lanceolate, apex usually coarsely toothed, plants often brick red in color  Bryoerythrophyllum recurvirostre 11. Leaves oblong-ligulate, apex never toothed; plants dark green to nearly black, never red  Didymodon  128  78. Leaf margins always bistratose, lamina occasionally bistratose i n spots; always on soil  Trichostomopsis australasiae 78. Leaf margins unistratose, rarely bistratose near apex; plants on rock or soil  79.  79. Median leaf cells with several low papillae per cell or appearing smooth; peristome teeth long and twisted; on soil Barbula unguiculata 79. Median leaf cells with several simple papillae per cell; peristome teeth simple, slender, erect, not twisted; on soil or rock  Didymodon  80. Leaves complanate, arranged in two rows or appearing so  81.  80. Leaves erect, spreading, not complanate  83.  81. Costa strong, to the middle o f the leaf or beyond; plants on wet rock or moist soil  Porotrichum 81. Costa short, double or lacking; plants on soil or rotten logs  78.  82. Leaf margin serrate to serrulate in the upper half; numerous branch like Propagula occur in the leaf axils, abundant in the spring; alar cells not differentiated  Pseudotaxiphyllum elegans  82. Leaf margin entire; propagula lacking; alar cells inflated, auriculate and decurrent  Plagiothecium 83. Leaves keeled, conduplicate, median cells long and narrow; plants usually forming long "stringy" mats; always in wet places Fontinalis 83. Leaves not keeled, conduplicate, median cells variable, never forming "stringy" mats; on rock, soil or trees, in wet or dry places 84. 84. Median leaf cells papillose with one or more papillae per cell or by projecting cell ends .... 85. 84. Median leaf cells not papillose  91.  85. Costa short, double, ending before the middle o f the leaf or lacking  86.  85. Costa double or single, extending to the mid-leaf or beyond  88.  86. Leaves strongly julaceous; median leaf cells papillose by projecting cell ends; alar cells strongly differentiated, extending up the margins; peristome teeth present; on rocks and trees Pterogonium gracile  129  86. Leaves never strongly julaceous, lacking a costa; median leaf cells with one or more papillae per cell; alar cells not differentiated; peristome lacking; on rocks  82.  87. Sporophyte immersed; perichaetial bracts with ciliate hairs on the margins; plants grayishgreen Hedwigia stellata 87. Sporophyte exserted; perichaetial bracts without ciliate hairs; plants reddish-brown  Pseudobraunia californica 88. Stems without paraphyllia; leaves crisped and contorted when dry; on soil, rocks or trees..  Claopodium 88. Stems with paraphyllia; leaves never crisped and contorted when dry  89.  89. Plants dendroid or frondiform; branch leaves coarsely serrate at the apex; on trees or occasionally rock Dendroalsia abietina 89. Plants never dendroid or frondiform; leaf margins plane or serrate, never coarsely serrate 90. 90. Plants with regular pinnate branching (sometimes irregularly) forming large mats on soil or rotten logs; median leaf cells long and narrow Hylocomium splendens 90. Plants branching + pinnately; forming small mats on rock; median leaf cells shorter  Heterocladium 91. Plants dendroid; stems matted with paraphyllia; on soil or rotten logs i n wet areas; (Leucolepis lacks paraphyllia)  Climacium dendroides  91. Plants not dendroid: stems with or without paraphyllia  92.  92. Costa short and double, never reaching beyond the middle o f the leaf, or lacking  93.  92. Costa long, extending to the middle o f the leaf or beyond  97.  93. Alar cells differentiated, inflated or quadrate  94.  93. A l a r cells scarcely differentiated  96.  94. Stems red; alar cells yellow-brown and thick-walled; on soil 94. Stems not red; alar cells hyaline, inflated or quadrate 95. Leaves straight, apex obtuse or rounded; on wet soil  130  Pleurozium schreberi 95. Calliergonella cuspidata  95. Leaves falcate to circinate, often ending in a filiform point; on wet or dry soil, rocks, trees and rotten logs Hypn urn 96. Costa wanting; plants filiform, forming thin mats on rock or soil in wet areas  Platydictya jungermannioides 96. Costa present; plants coarse, forming mats on soil, rotten logs or rocks....Rhytidiadelphus 97. Costa variable + with supplementary costae or double  98.  97. Costa always single and well developed, never having supplemental costae  101.  98. Stems with numerous paraphyllia  99.  98. Stems without paraphyllia  100.  99. Alar cells differentiated, extending up the leaf margins; branch leaves costa short or lacking; stems sometimes flagelliform; on trees or rock Alsia califomica 99. Alar cells quadrate to rectangular, not extending up the margins; costa double, to the middle o f the leaf; stems robust, never flagelliform; on soil Rhytidiopsis robusta 100. Alar cells forming a large triangular group, oval to transversely elongate; median leaf cells elongate; costa + supplemental costae; on trees or rocks Antitrichia 100. Alar cells not forming a large triangular group, median leaf cells thick and pitted; costa without supplemental costae; on soil, rotten logs and rarely rock Rhytidiadelphus 101. Leaves broadly ovate, suborbicular or broadly cordate-ovate  102.  101. Leaves ovate, ovate-lanceolate or lanceolate  107.  102. Stem and branch leaves julaceous  103.  102. Branch and stem leaves not julaceous  104.  103. Plants on rock in or near water; leaf apex obtuse 103. Plants on soil in lawns or old fields; leaf apex apiculate  Scleropodium obtusifolium Pseudoscleropodium  purum 104. Plants in wet locations; regularly or irregularly pinnate, branches spreading 104. Plants always on dry soil; regularly pinnate, branches in one plane  105.  Trachybryum  megaptilum  131  105. Alar cells abruptly enlarged and inflated; usually found in bogs, swamps or along lake margins Calliergon giganteum 105. Alar cells somewhat differentiated + inflated, never abruptly enlarged  106.  106. Alar cells quadrate; stem leaves broadly cordate-ovate, decurrent; on wet soil or woody debris Eurhynchium praelongum 106. Alar cells + inflated, stem leaves ovate to suborbicular, on rock, submerged in water  Platyhypnidium riparoides  107. Leaves deeply plicate  Homalothecium  107. Leaves not deeply plicate, sometimes lightly plicate  108.  108. Leaves falcate-secund; usually in wet areas, often submerged  109.  108. Leaves not falcate-secund; plants in dry or wet area, usually never submerged  111.  109. Alar cells inflated and sharply defined; leaves straight or slightly falcate  Cratoneuron filicinum  109. Alar cells undifferentiated, or quadrate; leaves always falcate 110. Leaves falcate to circinate, alar cells not extending to the costa  110. Drepanocladus  110. Leaves falcate-secund, alar cells quadrate often extending to the costa Sanionia uncinata 111. Alar cells quadrate, sporophyte usually inclined to horizontal and arcuate, not shrunken under the mouth when dry 112. 111. Alar cells short rectangular to quadrate-rectangular, + inflated; sporophyte inclined to horizontal, rarely upright, often shrunken under the mouth when dry 115. 112. Leaves broadly cordate-ovate, decurrent; margins serrate; stems branching irregularly to regularly pinnate; usually in dry places on soil, rotten logs and rocks Eurhynchium 112. Leaves never broadly cordate-ovate; margins entire or serrate in upper half only  113.  113. Alar cells thick-walled, quadrate, forming well defined area; upper leaf margin coarsely serrate; on rocks, trees and soil, never in wet areas Isothecium 113. Alar cells quadrate to rectangular, never forming distinct areas; leaf margin plane throughout, sometimes serrate at the apex  132  114.  114. Leaves often concave, never plicate; branches often julaceous; on the base o f trees, soil or sometimes soil over rock Scleropodium 114. Leaves never concave, + plicate (never deeply); branches never julaceous; on soil, rocks or in streams Brachythecium 115. Median cells long, 60-130 x 4-6  ; alar cells + inflated; usually in wet areas often  submerged 115. Median cells short, 25-65 x 6-11  116. ; alar cells not inflated, i n damp areas  116. Leaves straight, margins and apex entire 116. Leaves slightly curved, margins serrate at the apex  117.  Leptodictyum riparium Warnstorfia fluitans  117. Basal leaf margins toothed with sharp often recurved teeth; (median cells never papillose); usually in wet calcareous areas  Conardia compacta  117. Basal leaf margins never toothed, never on calcareous rock or soil  133  Amblystegium  K E Y S T O SPECIES  Amblystegium 1. Alar cells quadrate to transversely elongate, basal cells usually quadrate, leaves usually shorter, not wide-spreading A. serpens var. serpens 1. Alar cells few, rectangular, basal cells rectangular; leaves large, wide-spreading A. serpens \ar.juratzkanum  Amphidium 1. Leaf cells smooth, margins not recurved; autoicous, often with sporophytes  A. lapponicum  1. Leaf cells papillose with 8 or more papillae per cell; margins recurved; dioicous, sporophytes infrequent  A. californicum  Andreaea 1. Leaves ecostate  A. rupestris  1. Leaves costate  A. megistospora  Antitrichia 1. Costa single to the middle o f the leaf, occasionally faint supplemental costae near the leaf base; branches with strong imbricate leaves when dry; cell walls not pitted A. californica 1. Costa single with several distinct supplemental costae extending almost to the middle o f the leaf; branches with leaves somewhat divergent dry or wet; cell walls pitted A. curtipendula  Atrichum 1. Leaves broad, apex obtuse; several diagonal rows o f teeth on the dorsal surface; usually dioicous; common A. selwynii 1. Leaves narrower, apex acute; teeth on dorsal surface few or lacking; monoicous; uncommon A. undulatum  134  Aulacomnium 1. Alar cells inflated, sometimes brown, stems matted with rhizoids, gemmae leaf-like in irregular apical clusters on an attenuate shoot; on wet soil or in bogs A. palustre 1. Alar cells not inflated; rhizoids few; gemmae common, fusiform, clustered i n a sphere at the apex o f a leafless shoot; on rotten logs and soil A. androgynum  Brachythecium 1. Plants large, (leafy shoots > 1 m m wide); leaves ovate-lanceolate, deltoid-ovate to ovate lanceolate, plicate  2.  2. Plants light green to whitish -green; leaves ovate-lanceolate, apex long accuminate, lightly plicate; margins plane or slightly recurved at the base; alar cells quadrate; seta smooth; usually i n dry open sites on soil B. albicans 2. Plants dark green to yellow green; leaves deltoid ovate, to broadly ovate, lightly plicate, margins usually recurved, alar cells rounded or inflated; seta rough throughout; habitat various 3. 3. Leaves scarcely plicate, deltoid-ovate, abruptly acute, alar cells rounded and inflated, often forming long decurrent auricles; usually in wet areas B. rivulare 3. Leaves very plicate, broadly ovate to deltoid ovate, gradually acuminate; alar cells inflated forming a row across the bottom o f the leaf; seta rough or smooth, occurring in wet or dry areas  4.  4. Plants often stoloniform, forming loose open mats; leaf apex not twisted at the tip; found on dry forested sites B. asperrimum 4. Plants not stoloniform, occurring in dense or loose tufts; leaf apex often twisted at the tip; found i n wet places, sometimes submerged in water... B.frigidum 1. Plants small (leafy shoots < 1 m m wide); autoicous; leaves ovate-lanceolate to lanceolate, weakly plicate at base  5.  5. Seta rough throughout with large papillae  B. velutinum var. velutinum  5. Seta smooth or sometimes weakly papillose at base  B. velutinum var. venustum  135  Bryum 1. Plants grayish to whitish green, with clear to silver-white leaf tips  B. argenteum  1. Plants green, greenish-brown to reddish-brown  2.  2. Leaf shoots julaceous when dry, reddish-brown to purplish brown; leaf tip blunt  B. miniatum 2. Leaf shoots not julaceous when dry, green to brownish-green, sometimes tinged with red; leaf tip pointed 3. 3. Autoicous; leaves ovate-lanceolate + long accuminate; rhizoids few to numerous, leaf margins + revolute 4. 4. Leaves long acuminate, slightly contorted when dry, margins revolute; stems matted with rhizoids; rarely synoicous; cilia appendiculate B. pallescens 4. Leaves not long acuminate, margins occasionally lightly revolute; a few rhizoids at the base; cilia rudimentary or lacking B. uliginosum 3. Dioicous or sometimes synoicous; leaves ovate-lanceolate to oblong-ovate; rhizoids lacking to numerous; leaf margins + revolute, entire or denticulate 5. 5. Synoicous; costa excurrent as a + toothed point, brood-bodies absent; leaf margins revolute almost to the apex 6. 6. Stems matted with rhizoids; cilia lacking, rudimentary or present and appendiculate 7. C i l i a appendiculate; leaves ovate-lanceolate, acute  7.  B. lisae var. cuspidatum  7. C i l i a lacking or rudimentary; leaves ovate-lanceolate, long acuminate  B. ambylodon  5. Dioicous, occasionally synoicous; costa excurrent, percurrent or ending well below the apex, brood bodies present or absent, leaf margins ± revolute 8. 8. Leaves oblong-ovate, widest above the middle; filamentous propagula or rhizoidal tubers sometimes present 9. 9. Leaves decurrent; filamentous propagula often found i n the axils o f the leaves.... B.flaccidum 9. Leaves not decurrent; brown tubers occasionally found on rhizoids  B. capillare  8. Leaves ovate, ovate-oblong, never widest above the middle, or ovate-lanceolate; propagula may or may not be present 10.  136  10. Propagula never found; stem matted at the base with rhizoids; leaves ovate-lanceolate, long accuminate, widest at the base, upper leaf cells long, 6-7:1 B. caespiticium 10. Propagula often present; stems + matted with rhizoids; leaves oblong-lanceolate to obovateconcave, concave, median leaf cells short, 2-3:1, rare 4:1 11. 11. Propagula bulbiform; leaves ovate-oblong, ovate-lanceolate + concave, obtuse or acute, margins entire 12. 12. RJhizoids few, not papillose, leaves obtuse or broadly acute, growing in calcareous sites  B. gemmiparum 12. Rhizoids few, usually papillose, leaves ovate-lanceolate, concave; not usually found in calcareous sites B. dichotomum 11. Propagula filiform; or with tubers on rhizoids; leaves obovate-concave, ovate lanceolate... 13. 13. Filamentous propagula in leaf axils, leaves ovate-lanceolate, margin border unistratose or bistratose 14. 13. Rhizoids with brown tubers; leaves obovate-concave, acute, margin border unistratose  B. canariense 14. Leaf border unistratose; stem matted with dense reddish-brown rhizoids; leaves twisted and contorted when dry B. pseudotriquetrum  Claopodium 1. Plants branching irregularly; stem leaves without hyaline hairpoints; leaf cells with one large . papilla C. whippleanum 1. Plants usually pinnate; stem leaves with hyaline hairpoints; leaf cells with one or several papillae per cell  2.  2. Leaf cells with a single large papilla per cell  C. crispifolium  2. Leaf cells with two or more papillae per cell  C. bolanderi  Dicranella 1. Leaves oblong or obovate, squarrose from a sheathing base, margins unistratose, (bistratose in part), plane to incurved on damp soil D. schreberiana 137  1. Leaves lanceolate + sheathing base, not squarrose, margins unistratose or bistratose, plane, incurved or recurved in part 2. 2. Costa up to 1/3 width of leaf base; seta yellow or red, leaf margins unistratose or bistratose  3.  3. Seta yellow, darkening with age; leaf margins plane to incurved, unistratose; on road and trail soil banks D. heteromalla 3. Seta red; leaf margins plane to lightly recurved and bistratose; on clay  D. howei  2. Costa not up to 1/3 width of leaf base; seta red; leaf margins bistratose below the middle and recurved on one or both sides, on clay or wet soil D. pacifica  Dicranum 1. Leaf tips usually broken off, plants forming short "stiff tufts . 2 cm tall, leaves straight; median leaf cells not pitted, quadrate, rounded to shortly rectangular, on rotten logs and coniferous tree trunks D. tauricum 1. Leaf tips intact, not broken off, plants forming tall "soft" tufts > 3 cm tall, leaves usually falcate; median cells + pitted, lamina unistratose or bistratose found on a wide variety of substratum  2.  2. Median leaf cells strongly pitted and thick-walled, lamina unistratose, costa with dorsal "wings"; papillae absent on leaf cells D. scoparium 2. Median leaf cells not pitted, thin or thick-walled, lamina often bistratose on upper margins, costa without dorsal "wings"; papillae common on leaf cells D.fuscescens  Didymodon 1. Leaf apex obtuse to rounded, costa strong, ending before the apex; somewhat contorted when dry; peristome short and irregularly divided; often in wet calcareous areas D. tophaceous 1. Leaf apex acute, costa + strong, ending in or before the apex, lightly contorted to strongly crisped and contorted when dry; peristome teeth long or short and twisted; usually in dry calcareous sites  2.  2. Peristome teeth short and twisted; leaf cells smooth or nearly so, margins recurved below the middle, plane above, slightly contorted when dry D. rigidulus var. gracilis 138  2. Peristome teeth long and twisted; leaf cells papillose (rare smooth) with 1 or more papillae per cell 3. 3. Costa ending in or just below the apex, wide at the base in x-section, guide cells 5-7 at the base, 3-4 in the middle D.fallax 3. Costa ending in the apex, not wider at the base in x-section guide cells 4-5  4.  4. Leaves straight to curved, shorter (up to 2.5 m m long), margins recurved to above the middle D. vinealis var. vinealis 4. Leaves flexuous, longer (up to 5 m m long), margins recurved only at the base....!), vinealis  var. flaccidus  Ditrichum 1.. Autoicous, leaves linear-lanceolate, bistratose, apex serrate; seta yellow  D. montanum  1. Dioicous, leaves lanceolate-subulate, unistratose, bistratose near apex, apex entire or serrate; seta red 2. 2. Stems matted with rhizoids; costa excurrent 2. Stems not matted with rhizoids; costa percurrent to excurrent  D.flexicaule D. heteromallum  Drepanocladus 1. Costa extending beyond the middle, not excurrent, narrow at the base; leaf margins entire and plane D. aduncus 1. Costa excurrent, rarely percurrent, very thick and wide at the base; leaf margins lightly to sharply denticulate D. crassicostatus  139  Eurhynchium 1. Plants with coarse side branches, simple pinnate, usually "flattened" in one plane; widespread on a variety of substrata, usually in dry mostly forested areas E. oreganum 1. Plants with slender side branches, irregularly or often bipinnately branched; usually in wet locations on soil or woody material 2. 2. Stem leaves ovate, never squarrose; stem and branch leaves obtuse to + acute E. pulchellum 2. Stem leaves broadly cordate, squarrose; stem and branch leaves narrowly acute E. praelongum  Fissidens 1. Laminal cells bulging, in x-section mostly twice as deep as wide, often arranged in rows; limbidium on all lamina of the leaf and usually ending a few cells below apex; usually fruiting in spring; on soil F. limbatus 1. Laminal cells not bulging, not twice as deep as wide in x-section, not arranged in rows; limbidium ending in leaf apex; usually fruiting in fall; on rock  2.  2. Plants synoicous or gonioautoicous  F. bryoides var. bryoides  2. Plants rhizautoicous  F. bryoides var. viridulus  Fontinalis 1. Plants robust to very robust; stem leaves large, 4-8 mm long sometimes to 10 mm, 3 - 6.5 mm or occasionally to 8.5 mm wide F. antipyretica var. gigantea 1. Plants medium in size, slender to slightly robust; stem leaves smaller , not longer than 6 mm, not wider than 4 mm 2. 2. Plants medium to slightly robust; leaves somewhat divergent and swollen on branches; stem leaves 2 - 4 mm (rare 4.5 mm) wide F. antipyretica var. antipyretica 2. Plants slender to medium; leaves simply imbricated on branches; .75 - 2 mm wide, sometimes up to 3.5 mm ..F. antipyretica var. oregonesis  140  Funaria 1. Plants large (stems 1.6 - 6 cm tall) and branched; seta 1.6 - 6 cm long; annulus present and red; common and widespread, on soil F. hygrometrica 1. Plants smaller (stems up to 0.6 cm tall), not branched; seta 0.4 - 1.5 cm long; annulus lacking, uncommon, on soil F. muhlenbergii  Grimmia 1. Plants autoicous; seta curved to arcuate when moist; leaves unistratose, margins recurved below the middle, 2 - 3 stratose in the upper part, hairpoints long up to 1/2 the length o f the leaf; on rock and cement in dry exposed sites G. pulvinata 1. Plants dioicous; seta straight to + arcuate; leaves unistratose or bistratose, margins plane or recurved, hairpoints short or long 2. 2. Shoots stiff, coarse and + julaceous when dry; leaves not keeled, bistratose, margins plane to incurved, densely covered with spiny hairpoints G. laevigata 2. Shoots not stiff and coarse, never julaceous; leaves keeled, unistratose, margins plane or recurved, hairpoint muticous to long 3. 3. Plants strongly twisted and contorted when dry; hairpoint muticous or forming a short hyaline point; multicellular filamentous gemmae sometimes on the back o f the costa; leaf margins plane or recurved at the middle; on shaded rock outcrops and cliffs G. torquata 3. Plants not strongly twisted and contorted when dry, often lightly twisted; hairpoints smooth or nearly so, length up to 1.2 mm; margins strongly recurved only on one side, occasionally multicellular brown gemmae present on the leaves; on rock in forested areas....G. trichophylla  Heterocladium 1. Leaf and stem cells smooth; leaves squarrose; psudoparaphyllia few to inconspicuous H. procurrens 1. Leaf and stem cells papillose with 1 4 papillae per cell; leaves not squarrose; pseudoparaphyllia serrate and papillose H. macounii  141  Homaloth ecium 1. Alar cells numerous, quadrate and clear; leaf margins entire; occurring on rock or soil over rock H. pinnatifidum 1. Alar cells not numerous, short, irregularly quadrate to rounded; leaf margins entire or with sharp teeth; on soil, rocks or tree trunks 2. Plants found on trees, very rarely on rock  2. 3.  3. Plants slender, pale yellow and shiny, often pinnate; basal marginal cells with numerous sharp recurved teeth; plants usually in tight stoloniferous mats H. nuttallii 3. Plants robust, dark yellowish green and weakly glossy, rarely pinnate; basal marginal cells occasionally toothed with a few teeth, plants usually i n loose mats H.fulgescens 2. Plants on soil, soil over rock or on rock, very rarely on trees  4.  4. Leaves ovate-lanceolate, acuminate with a broad often serrate apex, strongly plicate, not concave at the base; alar cells short, irregularly quadrate to rounded H. aeneum 4. Leaves ovate-lanceolate, acute to acuminate, leaves plicate, very concave at the base; alar cells short, quadrate to rounded H. arenarium  Hypnum 1. Cortical cells small, thick-walled  2.  2. Plants forming small tightly appressed mats on logs and tree trunks; leaf base very rounded at the insertion H. circinale 2. Plants forming large loose mats on rocks and dry soil; leaf base not rounded at the insertion H. cupressiforme 1. Cortical cells large and hyaline  3.  3. Alar cells inflated, hyaline, strongly differentiated cells toward costa strongly pigmented; usually forming large mats in wetter habitats H. dieckii 3. Alar cells not inflated, not strongly differentiated; forming extensive mats on trees, cliffs in well-drained habitats H. subimponens  142  Isothecium 1. Leaves oblong-lanceolate, widest at or just below the middle; branches julaceous when dry; alar cells numerous, strongly differentiated quadrate to transversely elongate J . cristatum 1. Leaves ovate-lanceolate, widest just above the base; irregularly branching + flagellate or julaceous; alar cells short and quadrate, not numerous or strongly differentiated  /. myosuroides  Mnium 1. Leaf border bistratose to multistratose without stereid cells, median cells with thickened corners; peristome teeth yellowish brown, seta single M. marginatum 1. Leaf border multistratose with central streid cells, median cells without thickened coiners; peristome teeth dark red, multiple seta M. spinulosum  Neckera 1. Leaf apex long acuminate with sharp often recurved teeth; dioicous, sporophyte exserted  N. douglasii 1. Leaf apex blunt or rounded, without sharp teeth, sometime apiculate, or serrate; autoicous, sporophyte immersed N. pennata  Orthotrichum 1. Stomata superficial; dioicous or autoicous  2.  2. Always found on non-calcareous rock; autoicous; capsule immersed to emergent, occasionally slightly exserted O. rupestre 2. Found on trees, very rarely on rock, autoicous or dioicous, capsule immersed to exserted.... 3. 3. Plants dioicous; brood bodies often abundant; capsule immersed to emergent, rarely exserted..  O. lyellii  143  3. Plants autoicous; brood bodies absent; capsule immersed to exserted  4.  4. Sporophytes exserted, lightly ribbed and not constricted under the mouth when dry; 1- 3 large conical or forked papillae per cell O. speciosum 4. Sporophyte immersed to 1/2, emergent or shortly exserted  5.  5. Sporophyte smooth when dry, immersed; endostome o f 16 teeth, erose, stout and densely papillose O. striatum 5. Sporophyte strongly 8 ribbed when dry, 1/2 emergent to shortly exserted, endostome 8 (rare 16) teeth, not erose or stout, reticulate-papillose O. affine 1. Stomata immersed; always autoicous  6.  6. Plants growing on rock, often calcareous  7.  7. Leaf lamina i n upper 1/2 bistratose, (occasionally with unistrtose streaks), sporophyte immersed to 1/2 emergent, calyptra sparsely hairy 7. Leaf lamina unistratose, capsule exserted, calyptra hairy to nearly smooth  O. hallii O. anomalum  6. Plants growing on trees  8.  8. Leaf apex obtuse to broadly acute, rarely apiclute or serrulate tip, sporophyte 2/3 emergent to shortly-exserted; calyptra sparsely hairy or naked O. tenellum 8. Leaf apex acuminate to sharply acute, sporophyte exserted or 1/2 emergent; calyptra naked...9. 9. Sporophyte long exserted; peristome teeth pale yellow  O. consimile  9. Sporophyte exserted to 1/2 emergent, never long exserted; peristome teeth orange-red  O. pulchellum  Plagiothecium 1. Plants large, whitish-yellow-green, not glossy; leaves complantate, symmetric, very undulate; dioicous P. undulatum 1. Plants smaller, various shades o f green, always glossy; leaves complantate, symmetric or asymmetric, never undulate; autoicous, rarely dioicous 2. Leaves symmetric, with a long filiform apex up to 1/3 the length o f the leaf  144  2.  P. piliferum  2. Leaves asymmetric, apex not filiform  3.  3. Alar cells decurrent, composed o f + large bulging inflated cells; rarely dioicous  P. denticulatum 3. Alar cells decurrent, composed o f a narrow band o f rectangular cells; always autoicous  P. laetum  Pohlia 1. Leaves whitish-green to light green  2.  2. Leaves whitish-bluish green, opalescent or with a metallic luster; on rock cliffs in crevices and on ledges P. cruda 2. Leaves without metallic luster and not opalescent; in wet seepy places on soil or rock  P. wahlenbergii 1. Leaves o f various shades of green, never whitish  3.  3. Plants growing mixed in with Sphagnum, leaves small, acute and narrowly lanceolate  P. sphagnicola 3. Plants found in various habitats, wet or dry; leaves larger, ovate to ovate-lanceolate  4.  4. Leaves little decurrent, often twisted when dry; costa strong without spines on the back, antheridia in pairs at the base o f the upper leaves; propagula sometimes present  P. nutans  4. Leaves decurrent, not twisted when dry, costa not as strong, with spines on the back near the apex; antheridia in large disk-like heads P. longibracteata  Plagiomnium 1. Plants never having stoloniferous shoots; leaves obovate, slightly decurrent; marginal teeth sharp, extending almost to the base o f the leaf; on trees, rotten logs and sometimes soil  P. venustum 1. Plants usually with stoloniferous shoots; leaves various + decurrent; marginal teeth + sharp; usually on soil 2. 2. Plants dioicous; leaves oblong to elliptic, acuminate, concave and strongly decurrent, marginal teeth sharp and to the base o f the leaf; on soil P. insigne  145  2. Plants synoicous; leaves variously shaped, + decurrent, marginal teeth + sharp, extending almost to the base o f the leaf; on soil 3. 3. Leaves ovate to obovate or oblong, acute, strongly decurrent, marginal teeth sharp; on wet soil near water P. medium 3. Leaves elliptic, rounded at the base and apex, sometimes apiculate, slightly deccurent; marginal teeth blunt (rarely sharp); on soil or soil over rock i n dry areas P. rostratum  Philonotis 1. Plants small; upper leaf cells papillose on the upper ends; rhizoids not abundant; on welldrained soil P. capillaris 1. Plants robust; upper leaf cells papillose on the lower ends; stems with numerous brown rhizoids; on soil i n wet areas P.fontana  Pogonatum 1. Leaves strongly crisped and contorted when dry, leaf margins bistratose  P. contortum  1. Leaves not crisped and contorted when dry, leaf margins unistratose  P. urnigerum  Polytrichum 1. Leaves with incurved margins, forming "flaps" over the lamellae 2. Costa excurrent as a long toothed hyaline awn 2. Costa excurrent as a short toothed red awn or awn not conspicuous  2. P. piliferum 3.  3. Leaves with a short toothed awn, rhizoids only on or near the base o f the stem, usually i n dry places on soil P. juniperinum 3. Leaves without a conspicuous awn, stems matted with white rhizoids, only found in Sphagnum bogs P. strictum 1. Leaves with plane margins, never incurved  4.  146  4. Lamellae 4- 8 cells tall, apical cell notched or dish shaped  P. commune  4. Lamellae 3 - 5 cells tall, apical cell not notched, slightly larger than the rest o f the cells  P. formosum  Porotrichum 1. Leaf apex coarsely toothed; stems with paraphyllia or pseudoparaphyllia, usually in wet places  P. bigelovii 1. Leaf apex finely toothed, stems without paraphyllia; on the base o f trees or rock  P. vancouveriense  Rhizomnium 1. Leaves elongate-ovate or elliptic; narrowed at the base; stems without rhizoids or rarely a few; on rotten logs or rock R. glabrescens 1. Leaves obovate; stems matted with reddish-brown rhizoids the entire length o f the stem; on wet soil R- magnifolium  Rhytidiadelph us 1. Leaves without a costa or costa short and double, leaf cells smooth 2. Leaves squarrose recurved, not plicate, alar cells hardly differentiated  2. R. squarrosus  2. Leaves plicate, not strongly squarrose recurved, alar cells somewhat enlarged and well defined R- lore us 1. Leaves with a costa, double and strong to the middle, leaf cells papillose by projecting end walls R' triquetrus  Racomitrium 1. Leaves with apparent hairpoints  2.  2. Hairpoints hyaline, strongly papillose  3. 147  3. Papillae confined to hairpoint  R. lanuginosum  3. Papillae on most of leaf cells, including hairpoint  4.  4. Super-alar cells usually elongate, thin-walled, slightly papillose, hairpoint not strongly decurrent, lightly papillose in the upper part R. ericoides 4. Super-alar cells mostly short, thick-walled, strongly sinuous, hairpoints very decurrent, papillose or denticulate throughout R. elongatum 2. Hairpoints hyaline, smooth or denticulate  5.  5. Margins o f lamina bistratose, dorsal side of costa with bulging cells, hairpoint long or short  R. occidentale 5. Margins o f lamina unistratose, rarely with a few bistratose spots, hairpoint usually long and decurrent 6. 6. Upper lamina cells below the hairpoint small, short, not as long as wide, costa channeled, hairpoint bluntly decurrent, not usually forming long thin bands down the margins  R. heterostichum 6. Upper lamina cells below the hairpoint long, often 6 - 7 x as long as wide, costa deeply channeled, hairpoint very decurrent, forming thin lines down the margins R. lawtonae 1. Leaves without hairpoints or hairpoints short, often muticous  7.  7. Leaf apex blunt, obtuse, lacking a hairpoint  8.  8. Apex broad, usually blunt-denticulate, leaf cells papillose, apical leaf cells short, less than 3 : 1  R. aciculare  8. Apex blunt, not denticulate, leaf cells smooth, apical leaf cells elongate, greater than 3 : 1  R. fasciculare 7. Apex blunt, acute, muticous or rarely with a very short hairpoint  9.  9. Leaf cells papillose, apex acuminate, muticous or with a short decurrent weakly denticulate hairpoint; alar cells somewhat differentiated R. varium 9. Leaf cells smooth, acute to narrowly obtuse or muticous apex; alar cells not or scarcely differentiated R. pacificum  148  Schistidium 1. O n rock along the coast, often within the ocean spray zone, leaves stiff and slightly contorted when dry S. maritimum 1. O n rock, never i n the ocean spray zone, leaves never contorted when dry  2.  2. Leaf apex denticulate, mostly blunt or obtuse, usually on rock i n or near fresh water  S. rivulare 2. Leaf apex never denticulate, apex obtuse to acute, muticous + a hairpoint, on rock i n dry locations 3. 3. Leaves with hairpoints or muticous, margins recurved or revolute throughout, costa stout and channeled S. apocarpum 3. Leaves without hairpoints, rarely muticous, margins + recurved, costa stout, not channeled, sometimes denticulate on the dorsal surface S. agassizii  Scleropodium 1. Plants strongly julaceous, tumid branches, occurring i n or around streams, leaves broadly ovate and very concave S. obtusifolium 1. Plants + julaceous and tumid branches, occurring in well-drained sites, never i n or near water, leaves various  2.  2. Leaves ovate to ovate-lanceolate, apex acute without an apiculus; branches not julaceous, never tumid or turgid S. cespitans 2. Leaves ovate, oblong-ovate to broadly ovate-lanceolate, apex with a + recurved apiculus, branches julaceous, often tumid or turgid 3. 3. Branches often flattened somewhat; leaf apex with the apiculus not or rarely recurved; on wood or rock S. tourettii var. colpophyllum 3. Branches always terete; leaf apex with the apiculus never recurved; usually on mineral soil  S. tourettii var. tourettii  149  Sphagnum 1. Cortical cells o f stem and branches with spiral fibrils, branch leaves broad, rough at the back of the apex 2. 2. Plants pale red to purplish red, green cells central, not exposed on either surface  S. magellanicum 2. Plants green, never showing red, green cells isosceles-triangular, exposed on the inner surface  3.  3. Branch cells with numerous large rounded pores; chlorophyll cells with ridges on outer face....  :  S. henryense  3. Branch cells with fewer large elliptic pores; chlorophyll cells smooth on outer face  S. palustre 1. Cortical cells o f stems and branches without spiral fibrils, branch leaves narrower, apex not rough at back 4. 4. Branch leaves curved, stem and branch leaves with numerous commisural pores forming "bead like" rows, plants yellow-green to orange S. subsecundum 4. Branch leaves not curved, stem and branch leaves with few pores, never forming "bead like" rows, plants green, yellow-green to wine-red 5. Green cells exposed on the inner surface, plants reddish or brownish  6.  6. Capitulum flat on top, stem leaves oblong-lingulate, rounded at apex 7. Plants and stems rusty brown, capitulum branches straight, with erect leaves 7. Plants and stems reddish, capitulum branches upcurved, leaves subsecund  5.  7. S.fuscum S. rubellum  6. Capitulum "globose-rounded", stem leaves oblong to oblong-triangular, broadly acute  S. capillifolium 5. Green cells exposed on the outer surface, plants not red  8.  8. Divergent branch leaves very squarrose-spreading throughout; capitulum terminal bud large, margins o f branch leaves not wavy when dry S. squarrosum 8. Divergent branch leaves squarrose-spreading in capitulum only, capitulum terminal bud small, branch leaves with wavy margins when dry S. recurvum  150  Tortella 1. Leaves soft and strongly twisted and contorted when dry, margins wavy, costa excurrent as a + serrulate point T. tortuosa 1. Leaves rigid and not strongly twisted or contorted when dry, margins not wavy, costa strong, leaf tips frequently broken off T.fragilis  Tortula 1. Plants on trees, usually having numerous propagula, rarely without propagula  2.  2. Propagula multicellular, spherical or globouse, borne on the upper surface o f the leaf  3.  3. Leaves obovate-oblong to spathulate from a narrow base, apex obtuse, costa percurrent, never forming a hairpoint T. latifolia 3. Leaves obovate-oblong, occasionally oblong-lanceolate, apex broadly rounded; costa excurrent forming a smooth hairpoint often yellow-brown at the base T. papillosa 2. Propagula leaf-like gemmae borne on the top of the stem amongst the leaves, or lacking ..4. 4. Gemmae lacking; leaves with costa excurrent as a long smooth or lightly toothed hairpoint  T. laevipila var. laevipila 4. Gemmae leaf-like and costate, muticous or mucronate; leaves with costa excurrent as a short hairpoint or hairpoint absent T. laevipila var. meridionalis 1. Plants on soil, rocks or sometimes on trees, without propagula  5.  5. Leaf margins bordered by 1 -4 rows o f smaller yellowish cells; without long hairpoints ....6. 6. Plants autoicous; sporangia 4 - 6 m m long; leaves oblong-lanceolate to long-apatulate, costa excurrent as a short or long mucro; peristome teeth with a high basal membrane.... T. subulata 6. Plants dioicous; sporangia 1.5-2 m m long; leaves oblong to ligulate, costa percurrent, apex with a mucronate tip; peristome teeth with a l o w basal membrane T. amplexa 5. Leaf margins not bordered; with long, smooth or denticulate hairpoints  7.  7. Plants autoicous; leaves oblong to spatulate, costa strong, excurrent as a long smooth hairpoint; on calcareous rock or cement  T. muralis  7. Plants dioicous or synoicous; leaves oblong-lingulate, costa excurrent as a long denticulate hairpoint; on soil, rocks or trees 151  8. Plants synoicous, leaves recurved to just below the middle, costa strong, excurrent as a hyaline + toothed awn, usually on soil or soil over rock, sometimes on trees T. princeps 8. Plants dioicous, leaves recurved to the apex or just below it; costa strong, excurrent as a very toothed hairpoint usually red at the base 9. 9. Leaves densely papillose with one tall antleroid papilla per cell; on dry soil, never on trees  T. papillosissima 9. Leaves densely papillose with several low C-shaped papillae per cell; on dry soil or trees  T. ruralis  Ulota 1. Plants autoicous, often fruiting; gemmae lacking; on trees 2. Leaf apex bluntly acute, plants occurring as dense tufted cushions  2. U. obtusiuscula  2. Leaf apex acute, filiform, ending in a row o f several clear cells; plants prostrate, never forming dense cushions U. megalospora 1. Plants dioicous, rarely fruiting; abundant brown gemmae occurring on the leaf tips, on rock or trees only along the coast U. phyllantha  152  CHAPTER 8 Summary and Conclusions  The San Juan Islands contain a diverse moss flora within a small geographical area (445.5 square kilometers, 172 sq. miles) consisting o f 224 species and varieties, comprising 33 families and 97 genera. Four species were found as new for the State o f Washington with one o f these new for the United States. This flora represents 37.4 % o f what Lawton (1971) defined as the Pacific Northwest flora. Although 43 endemic Western North American taxa are present, none o f these are endemic just to the San Juan Islands. This flora is based on 6021 collections made over a four year period from 159 sites, and an extensive search o f historical records from herbaria throughout the region. The collection provides documentation o f the flora for future taxonomic studies, potential reevaluation, and is the foundation for a state wide moss checklist. A county distribution map for each o f the species was made, and when combined with the vouchers, provides a substantial database to monitor the loss o f species and habitat within the islands. Keys to the genera and species were designed for future use by both amateurs and bryologists and their construction helped to clarify species concepts that were not clear. It is hoped that these keys ultimately w i l l be expanded and refined to become useful for the entire state. Detailed ecological observations were made for each collection and included information on habitat and substratum. Six general habitats and twenty-six sub-habitats were distinguished in order to determine their correlations to the mosses. Other than the habitat and substratum information provided in a paper by Schofield (1976) little has been published on this topic with reference to the Pacific Northwest. Therefore this study supplements the British Columbia information, and appears to represent the first such study for mosses in Washington State.  153  Habitats showing the highest diversity included woodlands (60.2%), outcrops (58.4%), and wetlands with 57.5% o f the flora, with the majority o f the species occurring on rock (72.7%) or soil (63.3%). Because freshwater sources are not abundant within the islands, the aquatic substratum showed lowest diversity with only 4.0% o f the flora. Each moss collection was correlated with the geological units presented by Brandon et al. (1988) and a database was established for comparing the relationship o f the mosses to the geologic unit from which they were collected. The relationship between geological unit and species composition showed that the geological units with the greatest area in the islands correlated with the larger number o f species. This higher species composition is a result o f the greater variety o f rock material found in the larger units and is reflected i n the Constitution Formation (the largest formation covering 38.54% o f the area) having 83.9% o f the flora. One unit, the Nanaimo Group, covered only 4.84% o f the area, yet harbored 43.7% o f the moss flora. This probably is the result o f the large quantity o f conglomerate material that occurs in this formation which in turn, provides an excellent substratum for mosses to colonize. This flora was completed as a qualitative study, therefore species diversity is presented from a floristic viewpoint. The relative high species diversity found in the islands is the result o f a number o f factors. First the large number o f collections (6021) established a substantial database reflecting the six habitats and twenty six sub-habitats found within the islands. This database provided the information needed to interpret the geographical distributions and to compare the flora to the adjacent mainland and island floras. Although this database is quite large, additional species will certainly be added to it as more field work is completed. This is reflected in the number o f species found on Henry Island. Only half a day was spent on this island, resulting in a species list composed o f 28 species. Even with the addition of the five historical species the list still does not reflect what most likely w i l l be found on the island when more field work is  154  done. This same observation holds true for the islands that were not visited during the present study. The size o f the island also influences the number o f species. This is a result o f the number o f habitats available; the larger the island, the greater the number o f habitats available to species. Size alone is not the only limiting factor, the amount o f rainfall available and the degree o f disturbance (past or present) also influences species diversity. Orcas Island, the largest (14,761 hectares or 36,431 acres), clearly demonstrates this by harboring 85.7% o f the flora. San Juan Island, the second largest (14,382 hectares, or 35,448 acres) contained only 67.8% o f the flora. The higher percentage o f species on Orcas Island demonstrates the increased number o f niches found i n the M t . Constitution area resulting from the higher elevations, increased rainfall, and more freshwater sites. It also appears that less o f Orcas Island has undergone disturbance, while San Juan Island has a considerable amount o f agricultural land. Non-human disturbance also plays a role i n determining what species survive in or colonize, an area. This is clearly suggested by the influence o f bird droppings on Goose Island and by the loss o f species as a result o f trees blown down by winter storms, with a consequent loss o f habitat, on Orcas and Jones Islands. The establishment o f a complete inventory o f a region's flora is necessary before any phytogeographical interpretations can be made. It appears that this study presents the first phytogeographical interpretation o f past and present moss distribution patterns for Washington State. The bulk o f the flora is composed of circumboreal species that are derived from a once widespread Arcto-Tertiary flora. During the Pleistocene these islands were thoroughly glaciated and the present flora represents, therefore, species that have migrated back into the region predominantly from southern refugial sites, or less possibly from northern refugial areas. This recolonization in the islands may have begun as early as 13,500 yr. B . P . (Deither et al. 1995) but the bulk o f the flora appears to have returned 155  during the hypsithermal interval described by Deevey and Flint (1957). A cooling trend began about 2000 yr. B.P. that probably caused the southern element species to retreat southward, with fragments persisting only in those areas where favorable conditions also remained. The San Juan and adjacent islands can be interpreted as a "modern" refugium for southern mediterranean type climate species. It is hoped that, through continued protection o f these islands, this unique ecosystem w i l l continue to remain in a relatively undisturbed state, and as additional field work is completed, the recorded diversity o f the flora o f the islands w i l l be increased.  156  Literature cited Adams, R . P . 1983. Intraspecific Terpenoid Variation i n Juniperus scopulorum: Evidence for Pleistocene Refugia and Recolonization in western North America. Taxon 32: 30-46. Anderson, L . E . 1990. A checklist o f Sphagnum in North America north o f Mexico. The Bryologist93: 500-501. Anderson, L . E . , H . A . Crum and W . R . Buck. 1990. List o f mosses o f North America north o f Mexico. The Bryologist 93: 448-499. Atkinson, S. and F.Sharpe. 1985. W i l d Plants o f the San Juan Islands. The Mountaineers. Seattle, Washington. Axelrod, D.I. 1958. Evolution o f the Madro-Tertiary Geoflora. Botanical Review 27: 433-509. Axelrod, D.I. 1973. History o f the mediterranean ecosystem i n California. In Mediterranean Type Ecosystems, Origin and Structure, F. diCastre and H . Mooney (eds.) Springer-Verlag, N e w York, N e w York. Axelrod, D.I. 1975. Evolution and Biogeography o f Madrean-Tethyan Sclerophyll Vegetation. Annals Missouri Botanical Gardens. 62: 280-334. Barkman, J.J. 1963. A contribution to the taxonomy o f Tortula laevipila - T. pagorum - complex. Revue Bryologique et Lichenologique 32: 183-92. Barnosky, C . W . 1981. A Record o f Late Quaternary Vegetation from Davis Lake, Southern Puget Lowland, Washington. Quaternary Research 16: 221-239. Barnosky, C . W . 1984. Late Pleistocene and early Holocene environmental history o f southwestern Washington State, U . S . A . Canadian Journal o f Earth Science 21: 619-629. Barnosky, C . W . 1985. Late Quaternary Vegetation in the Southwestern Columbia Basin, Washington. Quaternary Research 23: 109-122. Bates, J.W. 1982. Quantitative approaches i n bryophyte ecology. In Bryophyte Ecology, A . J.E. Smith (ed.) p. 1-44. Chapman H a l l , N e w York, N e w York. Benson, K . R . 1986. The Young Naturalist's Society: From Chess to Natural History Collections. Pacific Northwest Quarterly 77: 82-93. Bergeron, T. 1944. O n some meteorological conditions for the dissemination o f spores, pollen etc., and a supposed wind transport o f Aloina Spores from the region o f Lower Yenisey to Southwest Finland i n July 1936. Svensk Botanisk Tidskrift 38: 269-292.  157  Brandon, M . T . , D . S . Cowan, J.E. Muller and J.A. Vance.1983. Field Trip Guidebook, Trip 5 Pre Tertiary Geology o f San Juan Islands, Washington and Southeast Vancouver Island, British Columbia. Geological Association o f Canada, Victoria Section, Victoria, British Columbia Brandon, M.T.and D . S . Cowan, J . A . Vance. 1988. The Late Cretaceous San Juan Thrust System, San Juan Islands, Washington. Geological Society o f America Special Paper 221 Boulder, Colorado. Buck, W . R . and D . H . Norris. 1996. Hedwigia stellata and H. detonsa (Hedwigiaceae) in North America. N o v a Hedwigia 62: 361-370. Burnham, J.C. ed. 1971. Science In America, Historical Selections. Holt, Rinehart and Winston, Inc. N e w York, N e w York. Chapman, J.D. 1952. The climate o f British Columbia, Department o f Geography, University o f British Columbia, Vancouver, British Columbia. Christy, J . A . 1984. Additions to the Moss Flora o f Oregon. The Bryologist 83: 355-358. Christy, J . A . and J . A . Lyford, D . Wagner. 1982. Checklist o f Oregon Mosses. The Bryologist 85 22-36. Crum, H . A . 1972. The geographic origins o f the mosses o f North America's eastern deciduous forest. Journal o f the Hattori Botanical Laboratory 35: 269-98. Crum, H . A . 1973. Mosses o f the Great Lakes Forest. Contributions from the University Herbarium, V o l . 10. A n n Arbor, Michigan. Crum, H . A . 1984. Sphagnopsida, Sphagnaceae. N e w Y o r k Botanical Garden Series II part II. N e w York, N e w York. Crum, H . A . 1988. A Focus on Peatlands and Peat Mosses. The University o f Michigan Press. A n n Arbor, Michigan. Crum, H . A . and L . E . Anderson. 1981. Mosses o f eastern North America, V o l . 1-2. Columbia University Press. N e w York, N e w York. Crundwell, A . C . and E . N y h o l m . 1977. Dicranella howei Ren. & Card, and its relationship to D. varia (Hedw.) Schimp. Lindbergia 4: 35-38. Danner, W . R . 1966. Limestone Resources o f Western Washington. Dept. o f Conservation. Division o f Mines and Geology Bulletin N o . 52. Olympia, Washington. Daubenmire, R. 1968. Plant Communities A Textbook o f Plant Ecology. Harper and R o w , N e w York, N e w York. Deevey, E.S. and R . F . Flint. 1957. Postglacial hypsithermal interval. Science 125: 182-184. 158  Dethier, D.P., F. Pessl Jr., R . F . Keuler, M . A . Balzarini and D . R . Pevear. 1995. Late Wisconsinan glaciomarine deposition and isostatic rebound, northern Puget Lowland. The Geological Society o f America Bulletin 107: 1288-1303. Dickson, J . H . 1967. Pseudoscleropodium purum (Limpr.) Fleisch. on St. Helena and Its Arrival on Tristan da Cunha. The Bryologist 70: 267-268. Dietrich, W . E . 1975. Surface Water Resources o f San Juan C o . In Geology and Water Resources of the San Juan Islands. R . H . Russell (ed.). Water Supply Bulletin 46. Department o f Ecology. Olympia, Washington. Easterbrook, D . J . 1969. Pleistocene Chronology o f the Puget Lowland and San Juan Islands, Washington. Geological Society o f American Bulletin 80: 2273-2286. Eckel, P . M . 1996. Synopsis o f the Mosses o f Wyoming. Great Basin Naturalist 56: 197-204. Easterbrook, D . J . 1969. Pleistocene Chronology o f the Puget Lowland and San Juan Islands. Geological Society o f America Bulletion 80: 2273-2286. Emig, W . H . 1918. Mosses as Rock Builders. The Bryologist 21: 24-27. Flowers, S. 1973. Mosses: Utah and the West. Brigham Young University Press. Provo, Utah. Flugel, E . 1982. Microfacies Analysis o f Limestone. Springer-Verlag, N e w York, N e w York. Foster, A . S . 1904. Hercules Club. American Botanist 9: 6-8. Franklin, J.F. and C . T . Dyrness. 1973. Natural Vegetation o f Oregon and Washington. Pacific Northwest Forest and Range Experimental Station, U . S . D . A . Forest Service, General Technical Report P N W - 8 . Portland, Oregon. Frisvoll, A . A . 1983. A taxonomic revision o f the Racomitrium Grimmiales). Gunneria41: 1-181.  canescens group (Bryophyta,  Frisvoll, A . A . 1988. A taxonomic revision o f the Racomitrium heterostichum group (Bryophyta, Grimmiales) i n N . and C . America, N . Africa, Europe and Asia. Gunneria 59: 1-289. Frye, T . C . 1933. J.W. Bailey. The Bryologist 36: 82-83. Gams, H . 1932. Bryo-cenology (Moss-Societies). In Manual o f Bryology. Fr. Verdoorn (ed.). The Hague, Martinus Nihoff, Netherlands. Godfrey, J . L . D . 1977. The Hepaticae and Anthocerotae o f southwestern British Columbia. Ph.D. Thesis, Department o f Botany, The University o f British Columbia, Vancouver, British Columbia. Good, R. 1931. A Theory o f Plant Geography. The N e w Phytologist 30: 149-171.  159  Grebe, V . C . 1917. Studien zur Biologie und Geographe der Laubmoose. Hedwigia 59: 1-208. Grout, A . J . 1928-1940. Moss Flora o f North America North o f M e x i c o I - I V . Published by the Author. Newfane, Vermont. Hanson, H . P . 1943. A Pollen Study o f T w o Bogs on Orcas Island o f the San Juan Islands, Washington. Bulletin o f the Torrey Botanical Club 70: 236-243. Hayner, N . S . 1929. Ecological Succession in the San Juan Islands. In The Rural Community. E . W . Burgess (ed.) Papers and Proceedings o f the 23rd Annual Meeting American Sociological Society. University o f Chicago Press. Chicago, Illionis. Hebda, R . J . 1983. Late-glacial and postglacial vegetation history at Bear Cove B o g , Northeast Vancouver Island, British Columbia. Canadian Journal o f Botany 61: 3172-3192. Herzog, T. 1926. Geographie der Moose. Fischer, Jena. Heusser, C . J . 1960. Regional Vegetation. In Late-Pleistocene Environments of North Pacific North America. American Geographical Society Special Publication 35. Heusser, C . J . 1983. Vegetational History o f the Northwestern United States Including Alaska. In Late Quaternary Envinroments o f the United States V o l . 1. H.A.Wright (ed.). University of Minnesota Press. Minneapolis, Minnesota. Heusser, L . E . 1983. Palynology and Paleoecology o f postglacial sediments in an anoxic basin, Saanich Inlet, British Columbia. Canadian Journal o f Earth Science 20: 873-885. Hibbert, D . M . 1979. Pollen analysis of Late Quaternary sediments from two lakes in the Southern Puget Lowland, Washington. M . S . Thesis, University o f Washington. Seattle, Washington. Higinbotham, B . 1985. The Mini-Plant W o r l d o f Mosses and Lichens. The 1986 San Juan Islands Almanac 12: 44-47. Hoisington, B . L . 1979. A study o f the Brachythecium asperrimum-frigidum species complex. M . S c . Thesis, The University o f British Columbia, Vancouver, British Columbia. Holzinger, J . M . 1905. T w o Changes o f Name. The Bryologist 4: 54. Holzinger, J . M . and T . C . Frye. 1921. Mosses o f the Bureau o f Soils K e l p Expedition to Alaska. Publications Puget Sound Biological Station 3: 23-64. Horton, D . G . 1982. A Revision of the Encalyptaceae (Musci), with Particular Reference to the North American Taxa, Part I. Journal o f the Hattori Botanical Laboratory 53: 365-418. Horton, D . G . 1983. A Revision o f the Encalyptaceae (Musci), with Particular Reference to the North American Taxa, Part II. Journal o f the Hattori Botanical Laboratory 54: 353-532. 160  Howard, G . E . 1963. Theodore Christian Frye (1869 - 1962). The Bryologist 66: 124-136. Ignatov, M . S . and O . M . Afonina (eds.) 1992. Check-list o f mosses of the former U S S R . Arctoa 1: 1-85. Ireland, R . R . 1969. Taxonomic studies on the genus Atrichum in North America. Canadian Journal of Botany 47: 353-368. Ireland, R . R . 1970. Rhacomitrium lawtonae, a N e w Moss Species from British Columbia and Washington. The Bryologist 73: 707-712. Ireland, R . R . 197'1. Atrichum. J n E . Lawton, Moss flora of the Pacific Northwest, p. 31-33. The Hattori Botanical Laboratory, Nichinan, Japan. Ireland, R . R . 1982. Moss Flora of the Maritime Provinces. National Museums o f Canada Publications in Botany N o . 13. Ottawa, Canada. Ireland, R . R . , and J.R. Spence. 1987. Rhacomitrium pacificum, a new moss species from western North America. Canadian Journal of Botany 65: 859-862. Jackson, C.I. 1985. Exploration as Science: Charles Wilkes and the U . S . exploring Expedition, 1838-42. American Scientist 73: 450-461. Janssens, J . A . 1983a. Past and Present Record o f Drepanocladus crassicostatus sp. nov. (Musci: Amblystegiaceae) and the status of D. trichophyllus in North America. The Bryologist 86: 44-53. Janssens, J . A . 1983b. Past A n d Extant Distribution of Drepanocladus in North America, With Notes On The Differentiation O f Fossil Fragments. Journal of the Hattori Botanical Laboratory 54: 251-298. Klotz, 0 . 1917. The History O f The Forty-Ninth Parallel Survey West O f The Rocky Mountains. The Geographical Review 3: 328-387. Koponen, T. 1974. A guide to the Mniaceae in Canada. Lindbergia 2: 160-184. Krajina, V . J . 1959. Bioclimatic Zones in British Columbia. University of British Columbia Botanical Series 1. Vancouver, British Columbia. Krajina, V . J . 1965. Biogeoclimatic zones and classification o f British Columbia. Ecology o f Western North America V o l . 1. University of British Columbia, Vancouver, British Columbia. Lawton, E . 1960. Pseudoscleropodium 237.  purum i n the Pacific Northwest. The Bryologist 63: 235-  Lawton, E . 1965. A revision of the genus Homalothecium the Torrey Botanical Club 92: 333-354. 1.61  i n western North America. Bulletin of  Lawton, E . 1971. Moss Flora of the Pacific Northwest. The Hattori Botanical Laboratory. Nichinan, Japan. Lewinsky-Haapasaari, J. and B . C . Tan. 1995. Orthotrichum hallii Sull. & Lesq. N e w to Asia. Harvard Papers in Botany 7: 1-6. Lincoln, R . J . and G . A . Boxshall, P.F. Clark. 1982. A Dictionary of Ecology, Evolution and Systematics. Cambridge University Press, London, England. Little Jr., E . L . 1971. Atlas of United States Trees V o l . 1. Conifers and Important Hardwoods. Miscellaneous Publications N o . 1146. U . S . Dept. of Agriculture Forest Service, Washington, D . C . Longton, R . E . 1980. Physiologial Ecology o f Mosses. In The Mosses of North America. R . J . Taylor and A . E . Leviton (eds.). Pacific Division American Asscoiation for the Advancement of Science. 74-113. San Francisco, California. Lyall, D . 1864. Dr. D . Lyall on the Botany of North-West America. Journal of the Proceedings of the Linnean Society of London. Botany 7: 124-145. Macfadyen, W . A . 1928. O n the Deposition of Calcareous Tufa in a Mountain Stream at Binn, Canton Valais, Switzerland. The Geological Magazine 65: 1-5. Manuel, M . G . 1974. A Revised Classification o f the Leucodontaceae and a Revision o f the Subfamily Alsioideae. The Bryologist 77: 531-550. Mathewes, R . W . 1973. A palynological study of the vegetation changes in the University Research Forest, southwestern British Columbia. Canadian Journal o f Botany 51: 20852103. Mathewes, R . W . 1985. Climate Change In Canada 5. Syllogeus 55: 397-422. Mathewes, R . W . , L . E . Heusser. 1981. A 12000 year palynological record of temperature and precipitation trends in southwest British Columbia. Canadian Journal o f Botany 59: 707710. Maunder, W . J . 1968. Synoptic Weather Patterns in the Pacific Northwest. Northwest Science 42: 80-88. McCleary, J . A . and V . V . Green. 1971. A Checklist of Idaho Mosses. The Bryologist 74: 175180. Mcintosh, T.T. 1986. The Bryophytes of the Semi-Arid Steppe o f South-Central British Columbia. Ph.D. Thesis, Department of Botany, The University o f British Columbia, Vancouver, British Columbia. McLaughlin, S.P. 1989. Natural floristic areas of the western United States. Journal of Biogeography 16: 239-248. 162  M c L e l l a n R . D . 1927. The Geology o f the San Juan Islands. University o f Washington Publications in Geology 2. Seattle, Washington. Miller, H . A . 1982. Bryophyte evolution and geography. Biological Journal o f the Linnaean Society 18: 145-96. Miller, N . G . and L . J . Howe Ambrose. 1976. Growth i n Culture o f Wind-blown Bryophye Gametophyte Fragments from Arctic Canada. The Bryologist 79: 55-63. Mitten, W . 1864. Bryologia o f the Survey o f the Forty-Ninthy Parallel o f Latitude. Journal o f the Proceedings o f the Linnean Society o f London (Botany) 8: 12-55. Mozino, J . M . 1970. Noticias de Nutka, A n Account o f Nootka Sound i n 1792. University o f Washington Press, Seattle, Washington. Murray, B . M . 1987. Andreae schofieldiana and A. megistospora, species novae, and Taxonomic Criteria for Sect. Nerviae (Andreaeopsida). The Bryologist 90: 15-26. Nagano, I. 1969. Comparative Studies o f Moss Vegetations Developing on the limestone, chert, and other rocks lying adjacent to each other in the Chichibu Mountain Area, Central Japan. Journal o f the Hattori Botanical Laboratory 32: 155-203. Newcomb, C F . 1923. ed. Menzies' Journal o f Vancouvers' Voyage, A p r i l to October, 1792. Archives o f British Columbia. M e m o i r N o . 5. Victoria, British Columbia. Noguchi, A . 1987-1994. Illustrated Moss Flora o f Japan parts 1-5. The Hattori Botanical Laboratory. Nichinan, Japan. Nyholm, E . 1954-1969. Illustrated Moss Flora o f Fennoscandia II M u s c i . Fascicles 1-6. G W K Gleerup, Lund, Sweden. Parihar, N . S . and G . B . Pant. 1975. Bryophytes as Rock Builders-Some Calcicole Mosses and Liverworts Associated with Travertine Formation at Sahasradhara, Dehr Dun. Current Science 44: 61-62. Pentecost, A . 1996. Moss growth and travertine deposition: the significance o f photosynthesis, evaporation and degassing o f carbon dioxide. Journal o f Bryology 19: 229-234. Phillips, E . L . 1966. Washington Climate for the Counties o f Clallam, Jefferson, Island, San Juan, Skagit, Snohomish, and Whatcom, Washington State. Washington State University, Pullman, Washington. Pielou, E . C . 1991 After the ice age: the return o f life to glaciated North America. The University of Chicago Press, Chicago, Illinois Pijl, van der, L . 1972. Principles o f Dispersal in Higher Plants. Springer-Verlag. N e w York, N e w York.  163  Piper, C . V . 1906. Flora o f the State o f Washington, Contributions from the United States National Herbarium 11. Smithsonian Istitution. United States Museum, Washington, D . C . Powers, H . A . 1964. Volcanic A s h from Mount Mazama (Crater Lake) and from Glacier Peak. Science 144: 1334-1336. Raven, P . H . 1977. The California Flora, Chapter 4. In Terrestrial Vegetation o f California. M . G . Barbour and J.Major (eds.). John Wiley and Sons. N e w York, N e w York. Richards, P . W . 1932. Ecology. In Manual o f Bryology. Fr. Verdoorn (ed.). The Hague, Martnus Nijhoff, Netherlands. 367-395. Richardson, D . H . S . 1985. The Biology o f Mosses. John Wiley and Sons Inc. A . Halsted Press Book, N e w York, N e w York. Rigg, G . B . 1925. Some Sphagnum Bogs o f the North Pacific Coast o f America. Ecology 6: 260279. Rigg, G . B . 1929. Notes on the History o f Botany in the State o f Washington. The Washington Historical Quarterly 22(3): 163-173. Rigg, G . B . 1958. Peat Resources o f Washington State. Bulletin N o . 44. State o f Washington Department o f Conservation. Division o f Mines and Geology. Olympia, Washington. Rigg, G . B . and C.T. Richardson. 1934. The Development o f Sphagnum Bogs in the San Juan Islands. American Journal o f Botany 21: 610-622. Robinson, H . 1970. A Revision o f the Moss Genus Trichostomopsis.  Phytologia 20: 184-191.  Russell, R . H . ed. 1975. Geology and Water Resources o f The San Juan Islands, San Juan Co., Washington. Water Supply Bulletin N o . 46. Dept. o f Ecology. Olympia, Washington. Sama-Wojcicki, A . M and Champion, Davis. 1983. Holocene Volcanism i n the Conterminous United States and the Role o f Silicic Volcanic A s h Layers in Correlation o f LatestPleistocene and Holocene Deposits. In Late Quaternary Envinroments o f the United States V o l . 2. H . A . Wright (ed.) University o f Minnesota Press, Mineapolis, Minnesota. Sayre, G . 1971. Cryptogame Exsiccatae - A n Annotated Bibliography o f Published Exsiccatae o f Algae, Lichens, Hepatics and M u s c i . Memoirs o f The N e w Y o r k Botanical Garden 19 (2) : 175-276. Sayre,G. 1975a. Cryptogame Exsiccatae - A n Annotated Bibliography o f Published Exsiccatae o f Algae, Lichens, Hepatics and M u s c i . Memoirs o f The N e w Y o r k Botanical Garden 19 (3) : 362-363. Sayre, G . 1975 b. Illustrations o f the Lost Hepatics o f the Wilkes Expeditions. The Bryologist 78: 204-205.  164  Schlots, F . E . , A.O.Ness, J.R. Rasmussen, C . J . M c M u r p h y , L . L . M a i n , R . J . Richards, W . A . Starr, and S . H . Krashevski. 1962. Soil Survey San Juan County Washington. U S . Dept. of Agriculture, Soil Conservation Service. U S . Government Printing Office, Washington D . C . 73 pp. Schofield, W . B . 1965. Correlations Between the Moss Floras o f Japan and British Columbia, Canada. Journal o f the Hattori Botanical Laboratory 28: 17-42. Schofield, W . B . 1969. Phytogeography of Northwestern North America: Bryophytes and Vascular Plants. Madrono 20: 155-207. Schofield, W . B . 1970. A N e w Species o f Dicranella Endemic to Western North America. Bryologist 73: 702-706. Schofield, W . B . 1972. Bryology in arctic and boreal North America and Greenland. Canadian Journal o f Botany 50: 1111-1133. Schofield, W . B . 1974. Bipolar Disjunctive Mosses In The Southern Hemisphere, With Particular Reference To N e w Zealand. Journal o f the Hattori Botanical Laboratory. 38: 13-32. Schofield, W . B . 1976. Bryophytes o f British Columbia III: habitat and distributional information for selected mosses. Syesis 9: 317-354. Schofield, W . B . 1980. Phytogeography o f the mosses of North America (North o f Mexico). In The mosses o f North America. R . A . Taylor and A . E . Leviton (eds.) Pacific Division. A A A S . San Francisco, California, p. 131-170. Schofield, W . B . 1984. Bryogeography of the Pacific Coast of North America. Journal o f the Hattori Botanical Laboratory 55: 35-43. Schofield, W . B . 1985. Heterocladium macounii i n North America. Monographs In Systematic Botany. Missouri Botanical Garden 11: 133-145. Schofield, W . B . 1988a. Bryogeography and the bryophytic characterizatioon o f biogeoclimatic zones o f British Columbia, Canada. Canadian Journal o f Botany 66: 2673-2686. Schofield, W . B . 1988b. Bryophyte disjunctions in the Northern Hemisphere: Europe and North America. Botanical Journal o f the Linnean Society 98: 211-224. Schofield, W . B . 1992a. Bryophyte distribution patterns. In Bryophytes and Lichens i n a Changing Environment. J.W. Bates and A . M . Farmer (eds.) Oxford Unversity Press, Oxford. 103-130. Schofield, W . B . 1992b. Some Common Mosses of British Columbia. Royal British Columbia Museum Handbook. Queen's Printer. Victoria, British Columbia. Schofield, W . B . 1994. Bryophytes o f Mediterranean Climates in British Columbia. Hikobia 11: 402-414. 165  Schofield,W.B. and H . A . Crum. 1972. Disjunctions in Bryophytes. Annals of The Missouri Botanical Garden 59: 174-202. Schnooberger, I. 1942. Distribution o f Tortulapapillosa  W i l s . The Bryologist 45: 24-27.  Schuster, R . M . 1983. Phytogeography of the Bryophyta. In N e w Manual of Bryology, R . M . Schuster (ed.). 463-626. Shacklette, H . T . 1965. Bryophytes Associated with Mineral Deposits and Solutions i n Alaska. United States Geological Survey Bulletin 1198-C. Washington D . C . 1-17. Sharp, A . J . , H . Crum and P . M . Eckel (eds.). 1994. The Moss Flora o f M e x i c o . Memoirs o f The N e w Y o r k Botanical Garden V . 69: 1113 pp. N e w York, N e w Y o r k . Shaw, J. 1982. Pohlia Hedw. (Musci) in North and Central America and the West Indies. Contributions to the University of Michigan Herbarium 15: 210-295. Slack, N . 1977. Species Diversity and Community Structure i n Bryophytes. N e w Y o r k State Studies. N e w Y o r k State Museum Bulletin 428. The University o f the State o f N e w Y o r k . The State Education Department, Albany, N e w York. Smith, A . M . 1917. The early history of The Bryologist and the Sullivant M o s s Society. The Bryologist 20: 1-8. Smith, A . J . E . 1978. The Moss Flora of Britain and Ireland. Cambridge University Press. Cambridge. Spence, J. 1988. Bryum Hedw. in Western North America. The Bryologist 91: 73-85. Steere, W . C . 1940. Tortula in North America North of M e x i c o . The Bryologist 43: 98-109. Steere, W . C . 1978. North American Muscology and Muscologists, a B r i e f History. N e w Y o r k Botanical Garden, Bronx, N e w York. Stott, P. 1981. Historical Plant Geography, A n Introduction. George A l l e n and U n w i n . London, England. Takhtajan, A . 1986. Floristic Regions of the World. Unversity o f California Press. Berkely, California. Taylor, A . 1919. Mosses as formers of Tufa and o f Floating Islands. The Bryologist 22: 38-9. Vitt, D . H . 1971. The Infrageneric Evolution, Phylogeny, and Taxonomy of the Genus Orthotrichum (Musci) in North America. N o v a Hedwigia. 21: 683-711. Vitt, D . H . 1973. A revision of the genus Orthotrichum in North America, north of M e x i c o . Bryophytorum Bibliotheca Band 1 V e r l a g von J. Cramer. Lehre, Germany.  166  Vitt, D . H . and M . Ostafichuk, I . M . Brodo. 1973. Foliicolous bryophytes and lichens o f Thuja plicata in Western British Columbia. Canadian Journal o f Botany 51: 571-580. Vitt, D . H . , S.R. Gradstein and Z . Iwatzuki. 1985. A W o r l d listing o f Herbaria: Collectors, Bryologists, and Current Research. Bryophytorum Bibliotheca. Verlag von J. Cramer. Lehre, Germany. Vitt, D . H . and W . R . Buck. 1992. K e y to the moss genera o f North America north o f Mexico. Verlag von J. Cramer, D-3301 Lehre, Germany. Waitt, R . B . and R . M . Thorson. 1983. The Cordilleran Ice Sheet in Washington, Idaho and Montana. In Late Quaternary Envinroments o f the U . S . V o l . 1. H . A . Wright (ed.), p. 53-70. Univ. o f Minnesota Press, Minneaoplis, Minnesota. Welch, W . H . 1960. A Monograph of the Fontinalaceae. The Hague. Martinus, Nijhoff, Netherlands. Wilkes, C . 1849. Narrative o f the United States Exploring Expedition V o l . I V . Lea and Blancad. Philidelphia, Pennsylvania Wolf, J . A . 1969. Neogene Floristic and Vegetational History o f the Pacific Northwest. Madrono 20: 83-109. Worley, I. 1972. The Bryo-Geography o f Southeastern Alaska. Ph.D. thesis, Department o f Botany, The University o f British Columbia. Vancouver, British Columbia. Young, F . D , ed. 1905. Dr. John Scouler's Journal o f A Voyage to North West America. Quarterly Oregon Historical Society 6: 159-205. Zander, R . H . 1978. N e w Combinations in Didymodon (Musci) and A K e y to the Taxa in North America North o f M e x i c o . Phytologia 41: 11-32. Zander, R . H . 1981. Didymodon (Pottiaceae) i n M e x i c o and California: Taxonomy and Nomenclature o f Discontinuous and Nondiscontinuous Taxa. Cryptogamie, Bryologique et Lichenologique 2: 379-422. Zanten, B . O . van. 1978. Experimental Studies on Trans-Oceanic Long-Range Dispersal o f Moss Spores in the Southern Hemisphere. Journal o f the Hattori Botanical Labratory 44: 455482. Zanten, B . O . van and T. Pocs. 1981. Distribution and Dispersal o f Bryophytes. In Advances in Bryology. W . Schultze-Motel (ed.). Verlag von J. Cramer, D-3301 Lehre, Germany.  167  Appendix A Historical records that were not relocated. HPSU = Portland State University, W W B = Western Washington University, W T U = University of Washington.  Genus & Species Fontinalis antipyretica Bryum argenteum Alsia californica Antitrichia californica Leptodictyum riparium Neckera douglasii Orthotrichum lyellii Rhytidiopsis robutsta Dendroalsia abietina Ceratodon purpureus Polytrichum piliferum Porotrichum vancouveriense Racomitrium ericoides Polytrichum piliferum Hygrohypnum bestii Roellia roellii Andreaea rupestris Bryum creberrimum Climacium dendroides Distichium capillaceum Encalypta vulgaris Fontinalis antipyretica F. antipyretica var. oregonensis Heterocladium macounii Philonotis fontana Plagiomnium rostratum Pohlia wahlenbergi Polytrichastrum alpinum Polytrichum commune Pseudotaxiphyllum elegans Racomitrium affine Rhytidiopsis robusta Sphagnum mendocinum Tayloria serrata Distichium capillaceum Mnium spinulosum Timmia austrica Tortula subulata Polytrichastrum alpinum Antitrichia curtipendula Mnium spinulosum Racomitrium varium Timmia austrica Brachythecium velutinum Scleropodium touretii var. colophyllum  Island Blakely Flattop Henry Henry Henry Henry Henry Lopez Matia Matia Matia Matia Matia Matia Orcas Orcas San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan San Juan Spieden Spieden Spieden Spieden Stuart Sucia Sucia Sucia Sucia Waldron Waldron  Collector F.L. Spaulding T.C. Frye A.S. Foster A.S. Foster A.S. Foster T.C. Frye T.C. Frye F. Richardson D. Durrwachter L. O'Flaherty D. Durrwachter D. Durrwachter D. Durwachter D. Durwachter C M . Roberts E. Peterson D. Mullen D. Mullen D. Mullen T.C. Frye D. Mullen D. Mullen D. Mullen T.C. Frye J. Fritz T.C. Frye ? A . Daugherty D. Mullen T.C. Frye T.C. Frye T.C. Frye T.C. Frye Mrs. Hartge C M . Roberts J.E. Kirkwood J.E. Kirkwood C M . Roberts A.S. Foster T.C. Frye T.C. Frye A . M . Daugherty A . M . Daugherty A.S. Foster A.S. Foster  Date  Location o f Record  6 July 1919 10 July 1921 11 July 1904 24 July 1904 13 July 1904 3 July 1905 3 July 1905 April 1983 18 Feb. 1962 18 Feb. 1962 18 Feb. 1962 18 Feb. 1962 18 Feb. 1962 18 Feb. 1962 7 July 1925 ?  WTU WTU HPSU WTU HPSU WTU WTU Per. comm. WWB WWB WWB WWB WWB WWB WTU WTU WWB WWB WWB WWT WWB WWB WWB WTU FHL WWB WWB WTU WWB WTU WTU WTU WTU WTU WTU WTU WTU WTU WTU WTU WTU WTU WTU HPSU HPSU  23 July 1925 ? 29 June 1925 1 July 1923 21 July 1925 July 1908 3 July 1925 3 July 1921 July 1908 6 July 1930 25 June 1925 15 July 1923 3 July 1925 3 July 1923 ? 31 July 1925 10 July 1908 ? 5 July 1925 17 July 1926 17 July 1926 5 July 1925 12 July 1904 10 July 1907 10 July 1907 1 July 1923 1 July 1925 ? 12 July 1912  Appendix B Data on collecting bag.  DATE  T  R  SEC  LOC  ELEVATION  LAT  SITE  forest  meadow  canyon  beach  lake  swamp  marsh  LONG ridge  pond  trail  valley river  roadside  bog stream  PARTIAL  FULL  FULL SUN  SHADE  SHADE  EXPOSURE: N S E W  TREE  trunk  LOG  recent-fallen  SOIL  branch  gravel  sand  root decaying  clay  MESIC spray  stump  rotten  litter  ROCK cave  leaf  humus  siliceous ledge  roof o f overhang  XERIC waterfall  seep  HYDRIC submerged  169  over rock basic crevice  moist standing  wet flowing  Appendix C Species and habitat relationships. Maritime  G & S alca amju amse amca amla anme adme adru anca ancu atse atun auan aupa baun bapo blac bral bras brfr brri brve brvn brre byam byar byca bycn bycp bydi byfl byge byli by mi bypa bypl byps byul cagi cacu cepu  M A R  X X  M S E  X X  Meadows & Ridges M 0 0 0 c G M R s s P s X  Disturbed Sites c D D E A D M I E  Outcrops 0 R Q  0 0  c  0 0 N X  Wetlands 0  s  N  W B 0  X X X  X X  X  X  X X  X X X  X  X  X  X X X  X  X  X  X  X  W L A  X X X X X X X X  X X X  X X X  X  X  X X  X X  X  X  W p  W  W  W  o  s  E  s  s  X X  M  T  X  X  X X X  X X X X X  X  X  X X  X X  X  X  X  X X X  X X X X  X  X  X X X  X  X  X  X  X  X X X  X  X X X  X  X  X  X  X  X X  X X  X  X  X  X  X  X  X X  X  X  X  X  X  X  X  X  X  X X X  X  X X  X X X  X X  X  X  X  X  X  X  X  X X X  X  X X  X  X X  X  X X  X  X  X  X X  X X  X X X X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  170  X X  X  X  X  X X  X X X  X  0 A 0  X X  X X  C D M w w w 0 o 0 X  X  X  X  Woodlands  X X  X X  X  X  X  X  X X X  X X X  X  X  X  X X  G & S clbo clcr clwh clde coco coca crfi crla cyje deab deob dipe dihe diho dipa disc dici dcfu dcsc dcta difa difl diri dito divi dtfl dthe dtmo drad drcr drpa enci enpr envu epto euve euor eupr eupu fiad fibr fivi figr fili five foan fogi  M A R  M  s E  M C  0  G  s s  o  M P  0  R  s  c  E M  D A I  D D E  0  X  X  R Q  X  X  o o c  0 0  o s  N  N  X X X  X X X  w B 0  w L A  w p  w  0  E  w  w  s s s M  X  T X  C w  D w  M w  0  0 X X X  0  0  0  X  X X  X  X  X  X  X  X X X X X X  X X X X X  X X X X  A  X X X X  X X  X  X  X X  X  X X  X X  X X X X X  X X  X X X  X X  X  X  X X  X X X  X X  X X X X  X X  X X X  X  X X  X X X  X X  X X  X  X  X  X  X X X  X  X X  X  X  X X X  X X X X X  X X X  X X X X  X X X  X X X  X  X X  X X X X  X X X X  X  X X  X X  X  X  X  X  X  X X  X  X  X  X  X X X  X  X X  X  X  X X X X  X X  X X  X X X X  X X X  X  X X  X X  X X  X  X X X  X X  X X  X X  X  X  X X  X X  X X  X  X X  X X X  X  X X  X  X  X  X X  X  X X  X  X  X  171  X X  X X  X  G &  S  foor fuhy fumu grla grpu grto grtr gyae hest hema hepr hoae hoar hofu honu hopi hysp hyre hyci hycu hydi hysu iscr ismy lepy leri leac meme mnma mnsp nedo nepe onwa oraf oran orco orha orly orpu orru orsp orst orte phcu phca phfo phpy  M A R  M  X  M  S C E s s  0 M P  0 R  X  X  X X  X  0 G  X  X  X X X  X  X  s  c E M  D A I  D D E  X  X  X  X  X  X  X  X  o R Q  0  o c  X  X  X  X  X X  X  X X X X  X X  X X X  X X X  X  X  X X X X  0  s  N  w w w w w w c D M B L p s s s w w w 0 A E M T o 0 0 o X  X  X X  0 0 N  X X X  X  X X X X X X X  X X X X X  X X X  X X  X X  X  X X  X  X  X X X  X X  X  X X  X  X  X  X  X X X  X  X  X  X  X  X  X  X X X X X X X  X X X  X  X  X  X  X  X X  0 A 0  X X X X X X  X X X X  X X X  X  X X  X X X  X  X X  X X  X  X X X X X X X X  X X X  X X X X  X X X X  X  X X  X X X X  X  X  X X X X  X  X X X  X  X X X  X X  X X X  X X  X X  X  X  X X X  X X  X  X  X X X X X X X X X X X  X X X  X X X X X X X X  X X  X  X X X  X X X  X  X  X X  X  X X X X X X X X  X X  X  X  X  X  X  X  X X  X X  X X X  X  X  X  X  X  X  X  X  X X  X  X X  X  X X  X X X X X X  X X X X X  X  X  X X X X  X  X X  172  G  M  M  M  0  &  A R  S E  C  G  S plin plme plro plve plde plla plpi plun  X  Plju plri plac plsc poco pour pocr polo ponu posp powa pyal pyco pofo pyju pypi pyst pobi pova psca pspu psel  X  Ptgr ptga raac rael raer rahe rala ralw raoc rapa rava rhgl rhma rhlo rhsq rhtr rhro  s s  0 M  P X  0  R  s  c  E M  D A I  D D E  O R Q  X  o o c  X  0 0  0  N  N  X X  X X  X  s  w B  w L A  w p  w  0  w  w  s s s  E  M  T  X  X  X X  X  X X X X X  X X X  X X  0  X  X  X  c  D w  M w  0 X X X X X X  0 X  0 X  X X X  X X X  w  X  X  X X X X  X  X X  X X  X  X  X X  X  X  X X X  X  X  X X  X  X X  X X  X  X  X  X X  X  X  X X  X X  X X  X  X X X  •  X  X  X X  X  X  X  X  X X X  X X  X  X X X X X X  X X  X  X X  X X X  X X X  X  X X X X X X X X X X X X X  X  X  X  X  X  X  X  X X  X X X  X  X X X X X X  X X  X X X X  X X  X X  X  X X  X  X X  X  X X  X  X X  X  X  X X X X  X  X  X X  X  X  X X X X X  X  X  X  X X X  X  X  X  X  X  X  X X  X X  X  X X  X  X  X  x  X  X  X  X  X  X  X X  X  X X  X  173  X X  X  0  A 0  G & S saun scag scap scma scri scce scco scob spto spca spfu sphe spina  M A R X X X X X X  M  s E  M  o  0  C  G  S  S  M P  0 R  s  c E M  D A I  X X  o  0  w  w  o s  B 0  L A X  0  D D  0 R  E  Q  c  N  X X  X  X X  X X X X X  0  X  N  X  X  toru tosu trme trey trau ulme ulob ulph  X X X X X X X X X X X X  X X  X  X X X  X  c  D  M  w  w  w  T  0  0  0  X  X  X  X X  X  X X  X  X X  X  X X X  X X X  X  X  X  X X X  X  X  X X  X  X X X  X X X X  X  X  X  X X  X X  X X  X  X X  X  X  X X  X X  X X  X X X  X X X  X X  X X  X  X  X  X  X X  X X  X X  X  X  X  X X  X  X X X X  0 A 0  X  X  1  wafl weco zyvi  X X X  X  X X  s  M  w s  X X  X  X  w  X  X X X  w s E  X  1 1  r  0  X  r  sppa spre spru spsq spsu tepe tiau tier toft toto toam tola tolt tome tomu topa topp topr  w p  X  X  X  X X  X X  174  X X  X X X  X X X X X X X  X X  X X  X X  X  X X  X X  X  X X  X X X  X  Appendix D Acronyms for genus and species, arranged alphabetically by acronym. Acronym adme adru alca amca amju amla amse anca ancu anme atse atun auan aupa bapo baun blac bra! bras brfr brre brri brve brvn byam byar bycp bycn byca bydi byfl byge byli by mi bypa bypl byps byul cacu cagi cepu clbo clcr clde clwh coca coco crfi crla  Genus and Species Andreaea megistospora Andreaea rupestris Alsia californica Amphidium californicum Amblystegium serpens var. juratzkanum Amphidium lapponicum Amblystegium serpens var. serpens Antitrichia californica Antitrichia curtipendula Anacolia menziesii Atrichum selwynii Atrichum undulatum Aulacomnium androgynum Aulacomnium palustre Bartramia pomiformis Barbula unguiculata Blindia acuta Brachythecium albicans Brachythecium asperrimum Brachythecium frigidum Bryoerythrophyllum recurvirostre Brachythecium rivulare Brachythecium velutinum var. velutinum Brachythecium velutinum var. venustum Bryum amblyodon Bryum argenteum Bryum capillare Bryum canariense Bryum caespiticium Bryum dichotomum Bryum flaccidum Bryum gemmiparum Bryum lisae var. cuspidatum Bryum miniatum Bryum p aliens Bryum pallescens Bryum pseudotriquetrum Bryum uliginosum Calliergonella cuspidata Calliergon giganteum Ceratodon purpureus Claopodium bolanderi Claopodium crispifolium Climacium dendroides Claopodium whippleanum Conscinodon calyptratus Conardia compacta Cratoneuron filicinum Crumia latifolia  175  cyje dcfu dcsc dcta deab deob dici difa difl dihe diho dipa dipe diri disc dito divi drad drcr drpa dtfl dthe dtmo enci enpr envu epto euor eupr eupu euve fiad fibr figr fili five fivi foan fogi foor fuhy fumu grla grpu grto grtr gyae hema hepr hest hoae hoar hofu honu  Cynodontium jenneri Dicranum fuscescens Dicranum scoparium Dicranum tauricum Dendroalsia abietina Desmatodon obtusifolius Dicranoweisia cirrata Didymodon fallax Didymodon vinealis var. flaccidus Dicranella heteromalla Dicranella howei Dicranella pacifica Dichodontium pellucidum Didymodon rigidulus var. gracilis Dicranella schreberiana Didymodon tophaceous Didymodon vinealis var. vinealis Drepanocladus aduncus Drepanocladus crassicostatus Dryptodon patens Ditrichum flexicaule Ditrichum heteromalum Ditrichum montanum Encalypta ciliata Encalypta procera Encalypta vulgaris Epipterygium tozeri Eurhynchium oreganum Eurhynchium praelongum Eurhynchium pulchellum Eucladium verticillatum Fissidens adianthoides Fissidens bryoides var. bryoides Fissidens grandifrons Fissidens limbatus Fissidens ventricosus Fissidens bryoides var. viridulus Fontinalis antipyretica var. antipyretica Fontinalis antipytretica var. gigantea Fontinalis antipyretica var. oregonensis Funaria hygrometrica Funaria muhlenbergii Grimmia laevigata Grimmia pulvinata Grimmia torquata Grimmia trichophylla Gymnostomum aeruginosum Heterocladium macounii Heterocladium procurrens Hedwigia stellata Homalothecium aeneum Homalothecium arenarium Homalothecium fulgescens Homalothecium nuttallii  176  hopi hyci hycu hydi hyre hysp hysu iscr ismy leac lepy leri meme mnma mnsp nedo nepe onwa oraf oran orco orha orly orpu orru orsp orst orte phca phcu phfo phpy plac plde plin Plju plla plme plpi plri plro plsc plun plve pobi poco pocr pofo polo ponu posp pour pova powa  Homalothecium pinnatifidum Hypnum circinale Hypnum cupressiforme Hypnum dieckii Hymenostylium recurvirostre Hylocomium splendens Hypnum subimpqnens Isothecium cristatum Isothecium myosuroides Leucolepis acanthoneuron Leptobryum pyriforme Leptodictyum riparium Metaneckera menziesii Mnium marginatum Mnium spinulosum Neckera douglasii Neckera pennata Onocophorus wahlenbergii Orthotrichum affine Orthotrichum anomalum Orthotrichum consimile Orthotrichum hallii Orthotrichum lyellii Orthotrichum pulchellum Orthotrichum rupestre Orthotrichum speciosum Orthotrichum striatum Orthotrichum tenellum Philonotis capillaris Phascum cuspidatum Philonotis fontana Physcomitrium pyriforme Pleuridium acuminatum Plagiothecium denticulatum Plagiomnium insigne Platydictya jungermannioides Plagiothecium laetum Plagiomnium medium Plagiothecium piliferum Platyhypnidium riparioides Plagiomnium rostratum Pleurozium schreberi Plagiothecium undulatum Plagiomnium venustum Porotrichum bigelovii Pogonatum contortum Pohlia cruda Polytrichum formosum Pohlia longibracteata Pholia nutans Pohlia sphagnicola Pogonatum urnigerum Porotrichum vancourveriense Pohlia wahlenbergii x  177  psca psel pspu ptga Ptgr pyal pyco PYJu pypi pyst raac rael raer rahe rala ralw raoc rapa rava rhgl rhlo rhma rhro rhsq rhtr saun scag scap scce scco scma scob scri spca spfu sphe spma sppa spre spru spsq spsu spto tepe tiau tier toam tofr toto tola tolt tome tomu topa  Pseudobraunia californica Pseudotaxiphyllum elegans Pseudoscleropodium purum Ptychomitrium gardneri Pterogonium gracile Polytrichastrum alpinum Polytrichum commune Polytrichum juniperinum Polytrichum piliferum Polytrichum strictum Racomitrium aciculare Racomitrium elongatum Racomitrium ericoides Racomitrium heterostichum Racomitrium lanuginosum Racomitrium lawtonae Racomitrium occidentale Racomitrium pacificum Racomitrium varium Rhizomnium glabrescens Rhytidiadelphus loreus Rhizomnium magnifolium Rhytidiopsis robusta Rhytidiadelphus squarrosus Rhytidiadelphus triquetrus Sanionia uncinatus Schistidium agassizii Schistidium apocarpum Scleropodium cespitans Scleropodium touretii var. colpophyllum Schistidium maritimum Scleropodium obtusifoliumSchistidium rivulare Sphagnum capillifolium Sphagnum fuscum Sphagnum henryense Sphagnum magellanicum Sphagnum palustre Sphagnum recurvum Sphagnum rubellum Sphagnum squarrosum Sphagnum subsecundum Scleropodium touretii var. touretii Tetraphis pellucida Timmia austriaca Timmiella crassinervis Tortula amplexa Tortella fragilis Tortella tortuosa Tortula laevipila Tortula latifolia Tortula laevipila var. meridionalis Tortula muralis Tortula papillosa  178  topp topr toru tosu trau trey trme ulme ulob ulph wafl weco zyvi  Tortula papillosissima Tortula princeps Tortula ruralis Tortula subulata Trichostomopsis australasiae Trichodon cylindricus Trachybryum megaptilum Ulota megalospora Ulota obusiuscula Ulota phyllantha Warnstorfia flu itans Weissia controversa Zygodon viridissimus var. rupestris  179  Appendix E Species relationships to substratum. A Q = aquatic, B L = beach logs, D W = decayed wood, F L = floating logs, EP = epiphytic, F O = folicolous, H D = humus and duff, PE = peatlands, RA = slate, RB = basalt, R C = conglomerate, R G = granite or acidic rock, R L = calcareous rock, and cement, RO = unknown rock, RS = sandstone, SC = soil clay, SD = soil dry, SW = soil wet. G E N U S A N D SPECEIS  SUBSTRATUM A  B Q L  Alsia californica Amblystegium serpens var. serpens A. serpens var. juratzkanum Amphidium californicum A. lapponicum Anacolia menziesii Andreaea megistospora A. rupestris Antitrichia californica A. curtipendula Atrichum selwynii A. undulatum Aulacomnium androgynum A. palustre Barbula unguiculata Bartramia pomiformis Blindia acuta Brachythecium albicans B. asperrimum B. frigidum B. rivulare B. velutinum var. velutinum B. velutinum var. venustum Bryoerythrophyllum recurvirostre Bryum amblyodon B. argenteum B. caespiticium B. canariense B. capillare B. dichotomum B. flaccidum B. gemmiparum B. lisae var. cuspidatum B. miniatum B. pallens B. pallescens B. pseudotriquetrum B. uliginosum Calliergon giganteum Calliergonella cuspidata  D W  E p  F L  X X X X  F  0  H D  P R R R R R E A B c G L  X X X X  X X X X X X  X X  X X  X  X X  X X  X  X  X X X  X  R  o s X X X  X  X X X X X X X X  R  s  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  s  s  c D w X X X X X  X X X X X X X X X X X X X X  X X X X X X X X X X X X X  X X X X X X X  X  X  X  X X X X X X X X X X X X X X X X X  X X X  X  180  X  X X  Genus & Species Ceratodon purpureus Claopodium bolanderi C. crispifolium C. whippleanum Climacium dendroides Conardia compacta Coscinodon calyptratus Cratoneuron filicinum Crumia latifolia Cynodontium jenneri Dendroalsia abietina Desmatodon obtusifolius Dichodontium pellucidum Dicranella heteromalla D. howei D. pacifica D. schreberiana Dicranoweisia cirrata Dicranum fuscescens D. scoparium D. tauricum Didymodon fallax D. rigidulus var. gracilis D. tophaceous D. vinealis var. vinealis D. vinealis var. flaccidus Ditrichum flexicaule D. heteromallum D. montanum Drepanocladus aduncus D. crassicostatus Dryptodon patens Encalypta ciliata E. procera E. vulgaris Epipterygium tozeri Eucladium verticillatum Eurhynchium oreganum E. praelongum E. pulchellum Fissidens adianthoides F. bryoides var. bryoides F. bryoides var. viridulus F. grandifrons F. limbatus F. ventricosus Fontinalis antipyretica var. antipyretica F. antipyretica var. gigantea F. antipyretica var. oregonensis  A Q  B L  E F F H p R R R R p L 0 D E A B c G X X X X X X X X X X X X X X X D W  R L  R  R  s s s  0 s c D w X X X X X X X X X X X X X  X  X X X  X X X  X X  X X X X  X  X X X X  X X X  X X X X X  X  X  X X X  X X  X  X  X X X X  X  X  X X X X  X X X  X  X X X X X X  X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  X X  X  X X  X X  X  X  X  X  X X  X X X X X X  Genus & Species Funaria hygrometrica F. muhlenbergii Grimmia laevigata G. pulvinata Grimmia torquata G. trichophylla Gymnostomum aeruginosum Hedwigia stellata Heterocladium macounii H. procurrens Homalothecium aeneum H. arenarium H. fulgescens H. nuttallii H. pinnatifidum Hylocomium splendens Hymenostylium recurvirostre Hypnum circinale H. cupressiforme H. dieckii H. subimponens Isothecium cristatum I. myosuroides Leptobryum pyriforme Leptodictyum riparium Leucolepis acanthoneuron Metaneckera menziesii Mnium marginatum M . spinulosum Neckera douglasii N . pennata Oncophorus wahlenbergii Orthotrichum affine 0 . anomalum O. consimile 0 . hallii 0 . lyellii 0 . pulchellum 0 . rupestre 0 . speceiosum 0 . striatum 0 . tenellum Phascum cuspidatum Philonotis capillaris P. fontana Physcomitrium pyriforme Plagiomnium inigne P. medium P. rostratum  A  Q  B L  D W  E  F L  p  F  0  H D  p E  R A  R B  R  c  X  X  X X X  X  X  X  X  X X X X X X X X X  X X  X X  X X X X X X X X  X X X X  X X X X X X  X X X X X  X X X X  X X  X X X X X  R G  R L  R  R  s s s  0 s c D X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  w X  X X X X  X X X X  X  X X X X X  X X X X X X X X X X X X X X  X  X X X X X X X X X  X  X X  X X X X  X X  X X X X  182  X X X X X X X  GENUS AND SPECIES  Plagiomnium venustum Plagiothecium denticulatum P. laetum P. piliferum P. undulatum Platydicrya jungermannioides Platyhypnidium riparioides Pleuridium acuminatum Pleurozium schreberi Pogoanatum contortum Pogonatum urnigerum Pohlia cruda P. longibracteata P. nutans P. sphagnicola P. wahlenbergii Polytrichastrum alpinum Polytrichum commune P. formosum P. juniperinum P. piliferum P. strictum Porotrichum bigelovii P. vancouveriense Pseudobraunia californica Pseudoscleropodium purum Pseudotaxiphyllum elegans Pterogonium gracile Ptychomitrium gardneri Racomitrium aciculare R. elongatum R. ericoides R. heterostichum R. lanuginosum R. lawtonae R. occidentale R. pacificum R. varium Rhizomnium glabrescens R. magnifolium Rhytidiadelphus loreus R. squarrosus R. triquetrus Rhytidiopsis robusta Sanionia uncinatus Schistidium agassizii S. apocarpum S. maritimum S. rivulare  B D E F F H p R R R R R R R s s s Q L W p L o D E A B c G L 0 s c D w A  X X X X X  X X X  X X  X  X X  X X X X X X X X X  X X X X X X X  X X  X  X  X X  X X X X X X X X X  X  X X  X X X  X X  X X X  X  X X X X X X  X X X  X X X  X X  X  X X  X  X X X X X X X X X X X X X X X X X X X X X  X X  X  X  X X  X X X  X X X  X  X X X X X  X X  X X X X  X X X X X X X X X X X X X X X  X  183  X  X X X X X X X X X X X X X X X  X  X  X  X X X X X  X X X  X X X X X X X X X X X X  X  Genus & Species Scleropodium cespitans S. obtusifolium S. touretii var. touretii S. touretii var. colpophyllum Sphagnum capillifolium S. fuscum S. henryense S. magellanicum S. palustre S. recurvum S. rubellum S. squarrosum S. subsecundum Tetraphis pellucida Timmia austriaca Timmia crassinervis Tortella fragilis T. tortuosa Tortula amplexa T. laevipila var. laevipila T. laevipila var. meridionalis T. latifolia T. muralis T. papillosa T. papillosissima T. princeps T. ruralis T. subulata Trachybryum megaptilum Trichodon cylindricus Trichostomopsis australasiae Ulota megalospora U . obtusiuscula U . phyllantha Warnstorfia fluitans Weissia controversa Zygodon viridissimus var. rupestris  A Q  B L  D W  F F H p R R R R R R R s s s p L o D E A B c G L 0 s c D w X x x X X E  X  X X  X  X X X X X X X X X X X X  X X X X  X X X  X X  X X X  X  X X X X X X X X X X X X X X X  X  X X X  X X  X X X X X X X X X X  X X X  X  X  X X X X X X X X X X X X X X  X X  X X X X X X X X X  184  X X X X X X X X X X  X X X X X X X X X X X X X X X X  Appendix F Species relationships to geologic unit. CF = Constitution Formation, C K = Chuckanut Formation, DB = Deadman Bay Volcanics, E G = East Sound Group, F C = Fidalgo Complex, L F = Lummi Formation, L S = Lopez Structural Complex, N G = Nanaimo Group, OC = Orcas Chert, QC = Quaternary Cover, SG = Speiden Group, T C = Turtleback Complex G E N U S & SPECIES Alsia californica Amblystegium serpens var. serpens A. serpens var. juratzkanum Amphidium californicum A. lapponicum Anacolia menziesii Andreaea megistospora A. rupestris Antitrichia californica A . curtipendula Atrichum selwynii A. undulatum Aulacomnium androgynum A. palustre Barbula unguiculata Bartramia pomiformis Blindia acuta Brachythecium albicans B. asperrimum B. frigidum B. rivulare B. velutinum var. velutinum B. velutinum var. venustum Bryoerythrophyllum recurvirostre Bryum amblyodon B. argenteum B. caespiticium B. canariense B. capillare B. dichotomum B. flaccidum B. gemmiparum B. lisae var. cuspidatum B. miniatum B. pallens B. pallescens B. pseudotriquetrum B. uliginosum Calliergon giganteum Calliergonella cuspidata Ceratodon purpureus Claopodium bolanderi C. crispifolium  C F X X X X X X X X X X X X X X  c  D E F L L K B G c F s X X X X X X X X  X  X X  X X X  X X X X X X  X X X X X X X  N 0 Q s T G c c G c X X X X X X  X  X X X X X X X X X X X X X X X X X X X X X X X  X X X X X X X X X X X  X  X X  X X X  X X X X X X X X X X X X X X X X  X X X X X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  X  X X X X X  X X  X X X X X X X X  X X X  185  X X  X X X X  X X  X X X X X X X X X  X X X X X X X X X X X X X X X X X X X X X X  Genus & Species Claopodium whippleanum Climacium dendroides Conardia compacta Coscinodon calyptratus Cratoneuron filicinum Crumia latifolia Cynodontium jenneri Dendroalsia abietina Desmatodon obtusifolius Dichodontium pellucidum Dicranella heteromalla D. howei D. pacifica D. schreberiana Dicranoweisia cirrata Dicranum fuscescens D. scoparium D. tauricum Didymodon fallax D. rigidulus var. gracilis D. tophaceous D. vinealis var. vinealis D. vinealis var. flaccidus Ditrichum flexicaule D. heteromallum D. montanum Drepanocladus aduncus D. crassicostatus Dryptodon patens Encalypta ciliata E. procera E. vulgaris Epipterygium tozeri Eucladium verticillatum Eurhynchium oreganum E. praelongum E. pulchellum Fissidens adianthoides F. bryoides var. bryoides F. bryoides var. viridulus F. grandifrons F. limbatus F. ventricosus Fontinalis antipyretica var. antipyretica F. antipyretica var. gigantea F. antipyretica var. oregonensis F. hygrometrica F. muhlenbergii Grimmia laevigata G. pulvinata G. torquata  C C D E F K B G X X X X X X X X X X X X X X X X X X X X X X  X X X X X X X X X  X X X X X X X X X X X X X X X X X X  F L L N 0 Q s T C F s G C c G c X X X  X  X X  X X X  X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  X X  X X X  X  X X X  X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 186  Genus & Species Grimmia trichophylla Gymnostomum aeruginosum Hedwigia stellata Heterocladium macounii H. procurrens Homalothecium aeneum H. arenarium H. fulgescens H. nuttallii H. pinnatifidum Hylocomium splendens Hymenostylium recurvirostre Hypnum circinale H. cupressiforme H. dieckii H. subimponens Isothecium cristatum I. myosuroides Leptobryum pyriforme Leptodictyum riparium Leucolepis acanthoneuron Metaneckera menziesii Mnium marginatum M . spinulosum Neckera douglasii N . pennata Oncophorus wahlenbergii Orthotrichum affine 0 . anomalum 0 . consimile 0 . hallii 0 . lyellii 0 . pulchellum 0 . rupestre 0 . speciosum 0 . sriatum 0 . tenellum Phascum cuspidatum Philonotis capillaris P. fontana Physcomitrium pyriforme Plagiomnium inigne P. medium Plagiomnium rostratum P. venustum Plagiothecium denticulatum P. laetum P. piliferum P. undulatum Platydictya jungermannioides Platyhypnidium riparioides  C c F K X X X X X X X X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X X  X  D E F B G c X X X X X X X X  X X X X X X X X X X X X X X X X  X X X X X X X X X X X X X X  X X X X X X  X X X X X X X X X X X X X X X X X X X  X X X X X X X  X X X X X X X X X X X X X  X X  X X X X X X X X X X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X X  X X X X X X X X X  X X X X X X X X X X X X X X X X  L L N 0 Q s T F s G c c G c X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  X X  187  X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  Genus & Species Pleuridium acuminatum Pleurozium schreberi Pogonatum contortum P. urnigerum Pohlia curda P. longibracteata P. nutans P. sphagnicola P. wahlenbergii Polytrichastrum alpinum Polytrichum commune P. formosum P. juniperinum P. piliferum P. strictum Porotrichum bigelovii P. vancouveriense Pseudobraunia californica Pseudoscleropodium purum Pseudotaxiphyllum elegans Pterogonium gracile Ptychomitrium gardneri Racomitrium aciculare R. elongatum R. ericoides R. heterostichum R. lanuginosum R. lawtonae R. occidentale R. pacificum R. varium Rhizomnium glabrescens R. magnifolium Rhytidiadelphus loreus R. squarrosus R. triquetrus Rhytidiopsis robusta Sanionia uncinatus Schistidium agassizii S. apocarpum Schistidium maritimum S. rivulare Scleropodium cespitans S. obtusifolium S. touretii var. touretii S. touretii var. colpophyllum Sphagnum capillifolium S. fuscum S. henryense S. magellanicum S. palustre  C c D E F L L N 0 Q s T F K B G c F s G c c G c X X X X X X X  X X  X X X X  X X X  X X  X X X  X X X  X X  X  X  X X X X X  X X  X X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  X  188  Genus and Species Sphagnum recurvum S. rubellum S. squarrosum S. subsecundum Tetraphis pellucida Timmia austriaca Timmiella crassinervis Tortella fragilis T. tortuosa Tortula amplexa T. laevipila var. laevipila T. laevipila var. meridionalis T. latifolia T. muralis T. papillosa T. papillosissima T. princeps T. ruralis T. subulata Trachybryum megaptilum Trichodon cylindricus Trichostomopsis australasiae Ulota megalospora U. obtusiuscula U . phyllantha Warnstorfia fluitans Weissia controversa Zygodon viridissimus var. rupestris  C c F K X X X X X X X X X X X  N 0 Q s T s G c c G c X X X X X X X X X X X X X X X X X X X X X X X X X X X  D E F L B G C F  L  X X  X  _x  X X  X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X  189  Appendix G Phytogeographic world distributions Legend: W N A = Endemic Western North America, M E D = Pacific North America - Mediterranean, C B = Circumboreal, C T = Circum-temperate, C P = Circumpolar, N P = North Pacific, B P = Bipolar, W W E = Western North America - Western Europe, W E N = Western North America - Eastern North America, W E A = Western North America - Eurasia, U N C = Unclassified, W D = Widespread, X = World Distribution, N = North America Distribution, A = Anthropogenic Introduction  G E N U S & SPECIES Alsia californica Amblystegium serpens var. serpens A. serpens var. juratzkanum Amphidium californicum A. lapponicum Anacolia menziesii Andreaea megistospora A. rupestris Antitrichia californica A. curtipendula Atrichum selwynii A. undulatum Aulacomnium androgynum A. palustre Barbula unguiculata Bartramia pomiformis Blindia acuta Brachythecium albicans B. asperrimum B. frigidum B. rivulare B. velutinum var. velutinum B. velutinum var. venustum Bryoerythrophyllum recurvirostre Bryum amblyodon B. argenteum B. caespiticium B. canariense B. capillare B. dichotomum B. flaccidum B. gemmiparum B. lisae var. cuspidatum B. miniatum B. pallens B. pallescens B. pseudotriquetrum  W N A X  M E D  C B  c T  c p  N p  B P  w w w E E  X X  N  W E A  U N C  w D  X  X X X X X  X  X X X X X X X X X X  X X X X  X X X X X  X X X  X  X  X  X X X X X  X  X X  X X  X X X  190  X  N  G E N U S & SPECIES Barbula uliginosum Calliergon giganteum Calliergonella cuspidata Ceratodon purpureus Claopodium bolanderi C. crispifolium C. whippleanum Climacium dendroides Conardia compacta Coscinodon calyptratus Cratoneuron filicinum Crumia latifolia Cynodontium jenneri Dendroalsia abietina Desmatodon obtusifolius Dichodontium pellucidum Dicranella heteromalla D. howei D. pacifica D. schreberiana Dicranoweisia cirrata Dicranum fuscescens D. scoparium D. tauricum Didymodon fallax D. rigidulus var. gracilis D. tophaceous D. vinealis var. vinealis D. vinealis var. flaccidus Ditrichum flexicaule D. heteromallum D. montanum Drepanocladus aduncus D. crassicostatus Dryptodon patens Encalypta ciliata E. procera E. vulgaris Epipterygium tozeri Eucladium verticillatum Eurhynchium oreganum E. praelongum E. pulchellum Fissidens adianthoides F. bryoides var. bryoides F. bryoides var. viridulus F. grandifrons  W N A  M E D  C B  C T  C P  N P  B P  w w E  w E N  W E A  U N C  w D  X X X  X X  X X X X X  X  X  X  X X X X X X X X X X X X X  X  X X X X X X X X  X X  X  X  X X N  X X X X  X X X  X X X X X X X  19  X X  G E N U S & SPECIES F. limbatus F. ventricosus Fontinalis antipyretica var. antipyretica F. antipyretica var. gigantea F. antipyretica var. oregonensis Funaria hygrometrica F. muhlenbergii Grimmia laevigata G. pulvinata G. torquata G. trichophylla Gymnostomum aeruginosum Hedwigia stellata Heterocladium macounii H. procurrens Homalothecium aeneum H. arenarium H. fulgescens H. nuttallii H. pinnatifidum Hylocomium splendens Hymenostylium recurvirostre Hypnum circinale H. cupressiforme H. dieckii H. subimponens Isothecium cristatum I. myosuroides Leptobryum pyriforme Leptodictyum riparium Leucolepis acanthoneuron Metaneckera menziesii Mnium marginatum M . spinulosum Neckera douglasii N . pennata Oncophorus wahlenbergii Orthotrichum affine 0 . anomalum 0 . consimile 0 . hallii 0 . lyellii 0 . pulchellum 0 . rupestre 0 . speciosum 0 . striatum O. tenellum  W N A  M E D  C B  C T  C p  N p  B P  w w E  w E N  W E A  U N C X  w D  X X X X X X X X  X X  N  X  X  X X X X  X X X X X X X  X X X  X  X X X X  X X X  X X  X X X X X N  X  X X X X  X X X X X  X  1  X X X  1  192  G E N U S & SPECIES Phascum cuspidatum Philonotis capillaris P. fontana Physcomitrium pyriforme Plagiomnium insigne P. medium P. rostratum P. venustum Plagiothecium denticulatum P. laetum P. piliferum P. undulatum Platydictya jungermannioides Platyhypnidium riparioides Pleuridium acuminatum Pleurozium schreberi Pogonatum contortum P. urnigerum Pohlia cruda P. longibracteata P. nutans P. sphagnicola P. wahlenbergii Polytrichastrum alpinum Polytrichum commune P. formosum P. juniperinum P. piliferum P. strictum Porotrichum bigelovii P. vancouveriense Pseudobraunia californica Pseudoscleropodium purum Pseudotaxiphyllum elegans Pterogonium gracile Ptychomitrium gardneri Racomitrium aciculare R. elongatum R. ericoides R. heterostichum R. lanuginosum R. lawtonae R. occidentale R. pacificum R. varium Rhizomnium glabrescens R. magnifolium  W N A  M E D  C B  c  c  T  p  N P  B P  w w w E E  X  N  W E A  U N C  w D  X  X X X  N  X X X X X  X X X  X X X X X  X X X X  X  X X X X X X X X X X  X X X X X X X X  X X X A X  X  N  X X X X X X X X X X X X X  193  X  G E N U S & SPECIES Rhytidiadelphus loreus R. squarrosus R. triquetrus Rhytidiopsis robusta Sanionia uncinata Schistidium agassizii S. apocarpum S. maritimum S. rivulare Scleropodium cespitans S. obtusifolium S. touretii var. touretii S. touretii var. colpophyllum Sphagnum capillifolium S. fuscum S. henryense S. magellanicum S. palustre Sphagnum recurvum S. rubellum S. squarrosum S. subsecundum Tetraphis pellucida Timmia austriaca Timmiella crassinervis Tortella fragilis T. tortuosa Tortula amplexa T. laevipila var. laevipila T. laevipila var. meridionalis T. latifolia T. muralis T. papillosa T. papillosissima T. princeps T. ruralis T. subulata Trachybryum megaptilum Trichodon cylindricus Trichostomopsis australasiae Ulota megalospora U. obtusiuscula U. phyllantha Warnstorfia fluitans Weissia controversa Zygodon viridissimus var. rupestris  W N A  M E D  C B  c T  c p  N P  B P  w w E  X X X  w E N N  W E A  U N C  w D  X X X X X X  X X X X X  N  X X X X X X X X X  X X X  X X X  X X  X X  X  X X  X X  X  X X X  X X X  X X  X  N  X X  X X  X X X  X X  X  X X X  X X  X  N X  X  194  N  Appendix H Phytogeographic distributions in the Pacific Northwest Legend: C = Coastal, CHI = Coastal - disjunct to Humid Interior, DIC = Dry Interior disjunct Coastal, R M C = Rocky Mountains - disjunct Coastal, M C = Mediterranean Climate, S A A = Subalpine- alpine, W = Widespread, A = Anthropogenic Introduction. M = Occurs in mediterranean climate along coast but in a few cases is disjunct. CHI  C  Genus & Species Alsia californica Amblystegium serpens var. serpens A. serpens var. juratzkanum Amphidium californicum A. lapponicum Anacolia menziesii Andreaea megistospora A. rupestris Antitrichia californica A . curtipendula Atrichum selwynii A . undulatum Aulacomnium androgynum A. palustre Barbula unguiculata Bartramia pomiformis Blindia acuta Brachythecium albicans B. asperrimum B. frigidum B. rivulare B. velutinum var. velutinum B. velutinum var. venustum Bryoerythrophyllum recurvirostre Bryum ambylodon B. argenteum B. caespiticium B. canariense B. capillare B. dichotomum B. flaccidum B. gemmiparum B. lisae var. cuspidatum B. miniatum B. pallens B. pallescens B. pseudotriquetrum B. uliginosum Calliergon giganteum Calliergonella cuspidata Ceratodon purpureus Claopodium bolanderi C. crispifolium  DIC  RMC  MC X  w X X  X X X X X M X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 195  A  C  Genus & Species Claopodium whippleanum Climacium dendroides Conardia compacta Coscinodon calyptratus Cratoneuron filicinum Crumia latifolia Cynodontium jenneri Dendroalsia abietina Desmatodon obtusifolius Dichodontium pellucidum Dicranella heteromalla D. howei D. pacifica D. schreberiana Dicranoweisia cirrata Dicranum fiiscescens D. scoparium D. tauricum Didymodon fallax D. rigidulus var. gracilis D. tophaceous D. vinealis var. vinealis D. vinealis var. flaccidus Ditrichum flexicaule D. heteromallum D. montanum Drepanocladus aduncus D. crassicostatus Dryptodon patens Encalypta ciliata E. procera E. vulgaris Epipterygium tozeri Eucladium verticillatum Eurhynchium oreganum E. praelongum E. pulchellum Fissidens adianthoides F. bryoides var. bryoides F. bryoides var. viridulus F. grandiffons F. limbatus F. ventricosus Fontinalis antipyretica var. antipyretica F. antipyretica var. gigantea F. antipyretica var. oregonensis Funaria hygrometrica F. muhlenbergii Grimmia laevigata G. pulvinata G. torquata G. trichophylla  CHI  DIC  RMC  MC X  w X X  X X X X M X X X X X X X X X M X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 196  A  C  Genus & Species Gymnostomum aeruginosum Hedwigia stellata Heterocladium macounii H. procurrens Homalothecium aeneum H. arenarium H. fulgescens H. nuttallii H. pinnatifidum Hylocomium splendens Hymenostylium recurvirostre Hypnum circinale H. cupressiforme H. dieckii H. subimponens Isothecium cristatum I. myosuroides Leptobryum pyriforme Leptodictyum riparium Leucolepis acanthoneuron Metaneckera menziesii Mnium marginatum M . spinulosum Neckera douglasii N . pennata Oncophorus wahlenbergii Orthotrichum affine 0 . anomalum O. consimile 0 . hallii 0 . lyellii 0 . pulchellum 0 . rupestre 0 . speciosum 0 . striatum 0 . tenellum Phascum cuspidatum Philonotis capillaris P. fontana Physcomitrium pyriforme Plagiomnium insigne P. medium P. rosiratum P. venustum Plagiothecium denticulatum P. laetum P. piliferum Plagiothecium undulatum Platydictya jungermannioides Platyhypnidium riparioides Pleuridium acuminatum Pleurozium schreberi  CHI  DCI  RMC  MC  W X  X X X X X X M X X X X X X X X X X X X M X X M X X X X M X M X X X X X X X X X X X X X X X X X X X X X 197  A  Genus & Species Pogonatum contortum P. urnigerum Pohlia cruda P. longibracteata P. nutans P. sphagnicola P. wahlenbergii Polytrichastrum alpinum Polytrichum commune P. formosum P. juniperinum P. piliferum P. strictum Porotrichum bigelovii P. vancouveriense Pseudobraunia californica Pseudoscleropodium purum Pseudotaxiphyllum elegans Pterogonium gracile Ptychomitrium gardneri Racomitrium aciculare R. elongatum R. ericoides R. heterostichum R. lanuginosum R. lawtonae R. occidentale R. pacificum R. varium Rhizomnium glabrescens R. magnifolium Rhytidiadelphus loreus R. squarrosus R. triquetrus Rhytidiopsis robusta Sanionia uncinata Schistidium agassizii S. apocarpum S. maritimum S. rivulare Scleropodium cespitans S. obtusifolium S. touretii var. touretii Scleropodium touretii var. colpophyllum Spahgnum capillifolium S. fuscum S. henryense S. magellanicum S. palustre Sphagnum recurvum S. rubellum S. squarrosum 198  C  CHI X  DIC  RMC  MC  w  A  X X X  X X  X X X X X X X X X X X  X X X X X X X X X X X X X X X X X X X X X X X X M X X X X X X X X X X X  C CHI DIC RMC MC w  Genus & Species Sphagnum subsecundum Tetraphis pellucida Timmia austriaca Timmiella crassinervis Tortella fragilis T. tortuosa Tortula amplexa T. laevipila var. laevipila T. laevipila var. meridionalis T. latifolia T. muralis T. papillosa T. papillosissima T. princeps T. ruralis T. subulata Trachybryum megaptilum Trichodon cylindricus Trichostomopsis australasiae Ulota megalospora U. obtusiuscula U. phyllantha Warnstorfia fluitans Weissia controversa Zygodon viridissimus var. rupestris  X X X X X X X X X X X X X X X X X X X X X X X X X  199  A  Appendix I Number of records and frequency of species on islands. Genus & Species Dicranum scoparium Polytrichum juniperinum Dicranoweisia cirrata Eurhynchium oreganum Grimmia trichophylla Rhytidiadelphus triquetrus Bryum capillare Isothecium myosuroides Polytrichum piliferum Schistidium maritimum Didymodon vinealis var. vinealis Ceratodon purpureus Tortula princeps Claopodium crispifolium Dicranum fuscescens Racomitrium ericoides Rhytidiadelphus loreus Dicranum tauricum Hedwigia stellata Homalothecium pinnatifidum Hylocomium splendens Plagiothecium undulatum Dicranella heteromalla Eurhynchium praelongum Isothecium cristatum Orthotrichum lyellii Timmiella crassinervis Ulota phyllantha Atrichum selwynii Leucolepis acanthoneuron Racomitrium elongatum Racomitrium heterostichum Zygodon viridissimus var. rupestris Homalothecium fulgescens Rhizomnium glabrescens Tortula rural is Antitrichia curtipendula Funaria hygrometrica Grimmia pulvinata Hypnum circinale Homalothecium nuttallii Metaneckera menziesii Orthotrichum pulchellum Plagiomnium insigne Plagiothecium laetum Aulacomnium androgynum Antitrichia californica  No. of Records 140 139 131 106 197 119 39 143 70 66 206 117 30 119 85 61 65 72 93 80 73 65 50 118 73 173 39 38 80 76 42 154 71 85 56 90 78 36 71 56 53 49 47 55 50 53 61 200  No. of Islands 27 27 26 26 26 26 23 23 23 23 22 21 21 20 20 19 19 18 18 18 18 18 17 17 17 17 17 17 16 16 16 16 16 15 15 15 14 14 14 14 13 13 13 13 13 12 11  % of Islands 96.4 96.4 92.8 92.8 92.8 92.8 82.1 82.1 82.1 82.1 78.5 75.0 75.0 71.4 71.4 67.8 67.8 64.2 64.2 64.2 64.2 64.2 60.7 60.7 60.7 60.7 60.7 60.7 57.1 57.1 57.1 57.1 57.0 53.5 53.5 53.5 50.0 50.0 50.0 50.0 46.4 46.4 46.4 46.4 46.4 42.8 39.2  Genus & Species Fissidens limbatus Hypnum subimponens Mnium spinulosum Orthotrichum consimile Anacolia menziesii Bryum pseudotriquetrum Neckera douglasii Schistidium apocarpum Ulota obtusiuscula Amphidium californicum Brachythecium albicans Bryum caespiticium Plagiothecium denticulatum Scleropodium touretii var. touretii Timmia austriaca Tortula muralis Bryum lisae var. cuspidatum Claopodium bolanderi Didymodon vinealis var. flaccidus Leptodictyum riparium Plagiomnium venustum Orthotrichum rupestre Pleurozium schreberi Pohlia cruda Porotrichum bigelovii Pseudotaxiphyllum elegans Trachybryum megaptilum Bartramia pomiformis Brachythecium velutinum var. velutinum Claopodium whippleanum Fissidens bryoides var. viridulus Hypnum dieckii Racomitrium lanuginosum Scleropodium cespitans Weissia controversa Alsia californica Amblystegium serpens var. serpens Calliergonella cuspidata Didymodon tophaceous Homalothecium arenarium LeDtobrvum pyriforme Pseudobraunia californica Tetraphis pellucida Brvum miniatum Brachythecium frigidum Bryum argenteum Dendroalsia abietina Homalothecium aeneum Hymenostylium recurvirostre Polytrichastrum alpinum Polvtrichum formosum Schistidium agassizii  No. of Records 35 57 36 67 48 44 55 53 24 75 17 34 35 26 42 26 15 44 17 37 36 35 28 18 33 25 24 34 16 11 6 26 14 10 16 14 10 15 15 5 9 8 25 9 23 6 12 11 10 22 5 6 201  No. of Islands 11 11 11 11 10 10 10 10 10 9 9 9 9 9 9 9 8 8 8 8 8 7 7 7 7 7 7 6 6 6 6 6 6 6 6 5 5 5 5 5 5 5 5 4 4 4 4 4 4 4 4 4  % of Islands 39.2 39.2 39.2 39.2 35.7 35.7 35.7 35.7 35.7 32.1 32.1 32.1 32.1 32.1 32.1 32.1 28.5 28.5 28.5 28.5 28.5 25.0 25.0 25.0 25.0 25.0 25.0 21.4 21.4 21.4 21.4 21.4 21.4 21.4 21.4 17.8 17.8 17.8 17.8 17.8 17.8 17.8 17.8 14.5 14.2 14.2 14.2 14.2 14.2 14.2 14.2 14 .2  Genus & Species Scleropodium obtusifolium Atrichum undulatum Brachythecium asperrimum Bryoerythrophyllum recurvirostre Bryum gemmiparum Bryum pallescens Cratoneuron filicinum Dichodontium pellucidum Dicranella schreberiana Ditrichum heteromallum Drepanocladus aduncus Encalypta ciliata Fissidens bryoides var. bryoides Grimmia torquata Gymnostomum aeruginosum Orthotrichum speciosum Orthotrichum striatum Pohlia nutans Ptychomitrium gardneri Racomitrium lawtonae Racomitrium occidentale Racomitrium pacificum Tortula amplexa Tortula laevipila var. meridionalis Tortula papillosissima Amblystegium serpens var. juratzkanum Aamphidium lapponicum Aulacomnium palustre Barbula unguiculata Crumia latifolia Dicranella howei Dicranella pacifica Didymodon rigidulus var. gracilis Encalypta procera Eucladium verticillatum Fissidens ventricosus Fontinalis antipyretica var. antipyretica Fontinalis antipyretica var. oregonensis Neckera pennata Orthotrichum affine Philonotis fontana Physcomitrium pyriforme Platydictya jungermannioides Pleuridium acuminatum Pohlia wahlenbergii Racomitrium varium Schistidium rivulare Scleropodium touretii var. colpophyllum Sphagnum capillifolium Sphagnum henryense Sphagnum recurvum Sphagnum squarrosum  No. of Records 15 7 4 6 3 7 11 11 4 11 3 5 11 11 5 6 3 14 8 4 7 4 3 8 4 3 3 16 2 7 2 4 3 3 6 5 7 9 2 3 6 2 3 4 4 14 4 2 3 12 10 9 202  No. of Islands 4 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2  % of Islands 14.2 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 10.7 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 "7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1  Genus & Species Tortella tortuosa Tortula laevipila var. laevipila Tortula papillosa Tortula subulata Trichodon cylindricus Trichostomopsis australasiae Ulota megalospora Warnstorfia fluitans Andreaea megistospora Andreaea rupestris Blindia acuta Brachythecium rivulare Brachythecium velutinum var. venustum Bryum amblyodon Bryum canariense Bryum dichotomum Bryum flaccidum Bryum pallens Bryum uliginosum Calliergon giganteum Climacium dendroides Conardia compacta Coscinodon calyptratus Cynodontium jenneri Desmatodon obtusifolius Didymodon fallax Ditrichum flexicaule Ditrichum montanum Drepanocladus crassicostatus Dryptodon patens Encalypta vulgaris Epipterygium tozeri Eurhynchium pulchellum Fissidens adianthoides Fissidens grandifrons Fontinalis antipyretica var. gigantea Funaria muhlenbergii Grimmia laevigata Heterocladium macounii Heterocladium procurrens Hypnum cupressiforme Mnium marginatum Oncophorus wahlenbergii Orthotrichum anomalum Orthotrichum hallii Orthotrichum tenellum Phascum cuspidatum Philonotis capillaris Plagiothecium medium Plagiothecium rostratum Plagiothecium piliferum Platyhypnidium riparioides  No. of Records 3 6 4 2 2 3 14 3 4 12 1 1 1 1 1 1 1 1 1 1 1 8 2 1 2 5 4 4 1 1 2 2 4 1 1 1 3 3 2 3 1 3 2 1 1 1 2 1 1 3  No. of Islands 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1  203  % of Islands 7.1 7.1 7.1 7.1 7.1 7.1 7.1 7.1 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5  Genus & Species Pogonatum contortum Pogonatum unigerum Pohlia longibracteata Pohlia sphagnicola Polytrichum commune Polytrichum strictum Porotrichum vancouveriense Pseudoscleropodium purum Pterogonium gracile Racomitrium aciculare Rhizomnium magnifolium Rhytidiadelphus squarrosus Rhytidiopsis robusta Sanionia uncinata Sphagnum fuscum Sphagnum magellanicum Sphagnum palustre Sphagnum rubellum Sphagnum subsecundum Tortella fragilis Tortula latifolia  No. of Record 1. 7 1 1 1 3 1 1 1 1 2 1 12 1 5 1 3 2 3 1 2  204  No. of Islands 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1  % of Islands 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5 3.5  Appendix J Species list arranged systematically.  Sphagnaceae Sphagnum capillifolium (Ehrh.) Hedw. S. fuscum (Schimp.) Klinggr. S. henryense Warnst. S. magellanicum Brid. 5. palustre L . s. recurvum P. Beauv. S. rubellum W i l s . S. squarrosum Crome S. subsecundum Nees in Sturm  Andreaeaceae Andreaea megistospora A. rupestris Hedw.  B . Murr.  Ditrichaceae Pleuridium acuminatum Lindb. Ditrichum flexicaule (Schwaegr.) Hamp. D. heteromallum (Hedw.) Britt. D. montanum Leib. Trichodon cylindricus (Hedw.) Schimp. Ceratodon purpureus (Hedw.) Brid.  Dicranaceae Dicranella heteromalla (Hedw.) Schimp. D. howei Ren. & Card. D. pacifica Schof. D. schreberiana (Hedw.) Hilf. ex C r u m & Anderson Oncophorus wahlenbergii Brid. Cynodontium jenneri (Schimp. i n Howie) Stirt. Dichodontium pellucidum (Hedw.) Schimp. Dicranoweisia cirrata (Hedw.) Lindb. ex M i l d e Dicranum fuscescens Turn. D. scoparium Hedw. D. tauricum Sapeh.  Fissidentaceae Fissidens adianthoides Hedw. F. bryoides Hedw. var. bryoides F. bryoides Hedw. var. viridulus (Sw.) Broth. F. grandifrons Brid. F. limbatus Sull. F. ventricosus Lesq.  Seligeriaceae Blindia acuta (Hedw.) Bruch & Schimp. in B . S . G .  205  Encalyptaceae Encalypta ciliata Hedw. E. procera Bruch E. vulgaris Hedw.  Pottiaceae Weissia controversa Hedw. Gymnostomum aeruginosum Sm. Hymenostylium recurvirostre (Hedw.) D i x . Eucladium verticillatum (Brid.) Bruch & Schimp. in B . S . G . Timmiella crassinervis (Hampe) L . K o c h Tortella fragilis (Hook. & W i l s in Drumm.) Limpr. T. tortuosa (Hedw.) Limpr. Trichostomopsis australasiae (Grev. & Hook.) Robins. Didymodon fallax (Hedw.) Zand. D. tophaceous (Brid.) Lisa D. vinealis (Brid.) Zand. var. vinealis D. vinealis (Brid.) Zand. var. flaccidus (Bruch & Schimp. in Schimp.) Zand. Bryoerythrophyllum recurvirsotre (Hedw.) Chen. Barbula unguiculata Hedw. Crumia latifolia (Kindb. in Mac.) Schof. Phascum cuspidatum Hedw. Desmatodon obtusifolius (Schwaegr.) Schimp. Tortula amplexa (Lesq.) Steere T. laevipila (Brid.) Schwaegr. var. laevipila T. laevipila (Brid.) Schwaegr. var. meridionalis (Schimp.) Wijk & Marg. T. latifolia Bruch ex Hartm. T. muralis Hedw. T. papillosa W i l s in Spruce T. papillosissima (Copp.) Broth. T. princeps De Not. T. ruralis (Hedw.) Gaertn. et al. T. subulata Hedw.  Grimmiaceae Coscinodon calyptratus (Hook, in Drumm.) C . Jens, ex Kindb. Dryptodon patens (Hedw.) Brid. Grimmia laevigata (Brid.) Brid. G. pulvinata (Hedw.) Sm. G. torquata Hornsch. in Grev. G. trichophylla Grev.  Ptychomitriaceae Ptycomitrium  gardneri  Lesq.  Funariaceae Physcomitrium pyriforme (Hedw.) Hampe Funaria hygrometrica Hedw. F. muhlenbergii Turn.  Bryaceae Pohlia cruda (Hedw.) Lindb. P. longibracteata Broth, in R o l l . 206  P. nutans (Hedw.) Lindb. P. sphagnicola (Bruch & Schimp.) Lindb. & A m e l l P. wahlenbergii (Web. & Mohr) Andrews Epipterygium tozeri (Grev.) Lindb. Leptobryum pyriforme (Hedw.) Wils. Bryum ambylodon C . M u l l . B. argenteum Hedw. B. caespiticium Hedw. B. canariense Brid. B. capillare Hedw. B. dichotomum Hedw. B. flaccidum Brid. B. gemmiparum De Not. B. lisae De Not. var. cuspidatum (Bruch & Schimp. in B . S . G . Marg. B. miniatum Lesq. B. pallens (Brid.) Sw. in Rohl. B. pallescens Schleich. ex Schwaegr. B. pseudotriquetrum (Hedw.) Gaertn. et al. B. uliginosum (Brid.) Bruch & Schimp. in B . S . G .  Mniaceae Mnium marginatum (With.) Brid. ex P. Beauv. M. spinulosum Bruch & Schimp. in B . S . G . Leucolepis acanthoneuron (Schwaegr.) Lindb. Rhizomnium glabrescens (Kindb.) T. K o p . R. magnifolium (Horik.) T. K o p . Plagiomnium insigne (Mitt.) T. K o p . P. medium (Bruch & Schimp. in B.S.G.) T. K o p . P. rostratum (Schrad.) T. K o p . P. venustum (Mitt.) T . K o p .  Aulacomniaceae Aulacomnium androgynum (Hedw.) Schwaegr. A. palustre (Hedw.) Schwaegr.  Bartramiaceae Anacolia mensiesii (Turn.) Par. Bartramia pomiformis Hedw. Philonotis capillaris Lindb. in Hartm. P. fontana (Hedw.) Brid.  Timmiaceae Timmia austriaca  Hedw.  Orthotrichaceae Zygodon viridissimus (Dicks.) Brid. var. rupestris Lindb. ex Hartm. Amphidium californicum (Hampe ex C . M u l l . ) Broth. A. lapponicum (Hedw.) Schimp. Orthotrichum affine Brid. O. anomalum Hedw. O. consimile Mitt. O. hallii Sull. & Lesq. in Sull. O. lyellii Hook. & Tayl. 207  O. pulchellum Brunt, in Winch. & Gateh. O. rupestre Schleich. ex Schwaegr. O. speciosum Nees in Sturm O. striatum Hedw. O. tenellum Bruch ex Brid. Ulota megalospora Vent, in Roll. U. obtusiuscula C. Mull. & Kindb. in Mac. & Kindb. U. phyllantha Brid. Fontinalaceae Foninalis antipyretica var. antipyretica Hedw. F. antipyretica Hedw. var. gigantea (Sull.) Sull. F. antipyretica Hedw. var. oregonensis Ren. & Card. Climaciaceae Climacium dendroides (Hedw.) Web. & Mohr. Anomodontaceae Pterogonium gracile (Hedw.) Sm. Hedwigiaceae Hedwigia stellata Hedenas Pseudobraunia californica (Lesq.) Broth. Leucodontaceae Alsia californica (Hook. & Arnott) Sull. Dendroalsia abietina (Hook.) Britt. Antitrichia californica Sull. in Lesq. A. curtipendula (Hedw.) Brid. Neckeraceae Neckera douglasii Hook. N. pennata Hedw. Metaneckera menziesii (Hook, in Drumm.) Steere Leskeaceae Claopodium bolanderi Best. C. crispifolium (Hook.) Ren. & Card. C. whippleanum (Sull. in Whipple & Ives) Ren. & Card. Pterigynandraceae Heterocladium macounii Best. H. procurrens (Mitt.) Jaeg. Thamnobryaceae Porotrichum bigelovii (Sull.) Kindb. P. vancouveriense (Kindb. in Mac.) Crum Amblystegiaceae Cratoneuron filicinum (Hedw.) Spruce Amblystegium serpens (Hedw.) Schimp. in B.S.G. var. serpens A. serpens (Hedw.) Schimp. in B.S.G. var. juratzkanum (Schimp.) Rau & Herv. Leptodictyum riparium (Hedw.) Warnst. Conardia compacta (C. Mull.) Robins. Sanionia uncinata (Hedw.) Loeske Drepanocladus aduncus (Hedw.) Warnst D, crassicostatus Janssens Warnstorfia fluitans (Hedw.) Loeske 208  Calliergon giganteum (Schimp.) Kindb. Calliergonella cuspidata (Hedw.) Loeske  Brachytheciaceae Homalothecium aeneum (Mitt.) Lawt. H. arenarium (Lesq.) Lawt. H. fulgescens (Mitt, ex C . M u l l . ) Lawt. H. nuttallii (Wils.) Jaeg. H. pinnatifidum (Sull. & Lesq.) Lawt. Trachybryum megaptilum (Sull.) Schof. Isothecium cristatum (Hampe) Robins. I. myosuroides Brid. Brachythecium albicans (Hedw.) Schimp. in B . S . G . B. asperrimum (Mitt.) Sull. B. frigidum ( C . M u l l . ) Besch. B. rivulare Schimp in B . S . G . B. velutinum (Hedw.) Schimp. in B . S . G . var. velutinum B. velutinum (Hedw.) Schimp. in B . S . G . var. venustum (De Not.) A r c . Scleropodium cespitans (C. Mull.) L . K o c h S. obtusifolium (Jaeg.) Kindb. in Mac. & Kindb. S. touretii (Brid.) L . K o c h var. touretii S. touretii (Brid.) L . K o c h var. colpophyllum (Sull.) Lawt. ex Crum Eurhynchium oreganum (Sull.) Jaeg. E. praelongum (Hedw.) Schimp. in B . S . G . E. pulchellum (Hedw.) Jenn. Platyhypnidium riparioides (Hedw.) D i x . Pseudoscleropodium purum (Hedw.) Fleisch. in Broth.  Plagiotheciaceae Plagiothecium denticulatum (Hedw.) Schimp. in B . S . G . P. laetum Schimp. in B . S . G . P. piliferum (Sw. ex Hartm.) Schimp. in B . S . G . P. undulatum (Hedw.) Schimp. in B . S . G .  Hypnaceae Platydictya jungermannioides (Brid.) Crum Hypnum circinale Hook. H. cupressiforme Hedw. H. dieckii Ren. & Card, in R o l l H. subimponens Lesq. Pseudotaxiphyllum elegans (Brid.) Iwats.  Hylocomiaceae Hylocomnium splendens (Hedw.) Schimp. in B . S . G . Rhytidiadelphus loreus (Hedw.) Warnst. R. squarrosus (Hedw.) Warnst. R. triquetrus (Hedw.) Warnst. Pleurozium schreberi (Brid.) Mitt. Rhytidiopsis robusta (Hook.) Broth.  Tetraphidaceae Tetraphis pellucida  Hedw.  209  Polytrichaceae Atrichum selwynii Aust. A. undulatum (Hedw.) P. Beauv. Pogonatum contortum (Brid.) Lesq. P. urnigerum (Hedw.) P. Beauv. Polytrichastrum alpinum (Hedw.) G . L . Sm. Polytrichum commune Hedw. P. formosum Hedw. P. juniperinum Hedw. P. piliferum Hedw. P. strictum Brid.  210  Appendix K Alphabetic list of Species, * = historical collection not relocated. Alsia californica (Hook. & Arnott) Sull. Amblystegium serpens (Hedw.) Schimp. i n B . S . G . var. serpens A. serpens (Hedw.) Schimp. in B . S . G . vw.juratzkanum (Schimp.) Rau & Herv. Amphidium californicum (Hampe ex C . M u l l . ) Broth. A. lapponicum (Hedw.) Schimp. Anacolia menziesii (Turn.) Par. Andreaea megistospora B . Murr. A. rupestris Hedw. Antitrichia californica Sull. in Lesq. A. curtipendula (Hedw.) Brid. Atrichum selwynii Aust. A. undulatum (Hedw.) P. Beauv. Aulacomnium androgynum (Hedw.) Schwaegr. A. palustre (Hedw.) Schwaegr. Barbula unguiculata Hedw. Bartramia pomiformis Hedw. Blindia acuta (Hedw.) Bruch & Schimp. i n B . S . G . Brachythecium albicans (Hedw.) Schimp. i n B . S . G . B. asperrimum (Mitt.) Sull. B. frigidum (C. M u l l . ) Besch. B. rivulare Schimp. in B . S . G . B. velutinum (Hedw.) Schimp. in B . S . G . var. velutinum B. velutinum (Hedw.) Schimp. in B . S . G . var. venustum (De Not.) A r c . Bryoerythrophyllum recurvirostre (Hedw.) Chen. Bryum ambylodon C . M u l l . B. argenteum Hedw. B. caespiticium Hedw. B. canariense Brid. B. capillare Hedw. B. dichotomum Hedw. B.flaccidum Brid. B. gemmiparum De Not. B. lisae De Not. var. cuspidatum (Bruch & Schimp. in B.S.G.) Marg. B. miniatum Lesq. B. pallens (Brid.) Sw. in Rohl. B. pallescens Schleich. ex Schwaegr. B. pseudotriquetrum (Hedw.) Gaertn. et al. B. uliginosum (Brid.) Bruch & Schimp. i n B . S . G . Calliergon giganteum (Schimp.) Kindb. Calliergonella cuspidata (Hedw.) Loeske Ceratodon purpureus (Hedw.) Brid. Claopodium bolanderi Best. C. crispifolium (Hook.) Ren. & Card. C. whippleanum (Sull. in Whipple & Ives) Ren. & Card. 211  Climacium dendroides (Hedw.) Web. & Mohr. Conardia compacta (C. M u l l . ) Robins. Coscinodon calyptratus (Hook, in Drumm.) C . Jens, ex Kindb. Cratoneuron filicinum (Hedw.) Spruce Crumia latifolia (Kindb. in Mac.) Schof. Cynodontium jenneri (Schimp. in Howie) Stirt. Dendroalsia abietina (Hook.) Britt. Desmatodon obtusifolius (Schwaegr.) Schimp. Dichodontium pellucidum (Hedw.) Schimp. Dicranella heteromalla (Hedw.) Schimp. D. howei Ren. & Card. D. pacifica Schof. D. schreberiana (Hedw.) Hilf. ex Crum & Anderson Dicranoweisia cirrata (Hedw.) Lindb. ex M i l d e Dicranum fuscescens Turn. D. scoparium Hedw. D. tauricum Sapeh. Didymodon fallax (Hedw.) Zand. D. rigidulus Hedw. var. gracilis (Schleich. ex Hook. & Grev.) Zand. D. tophaceous (Brid.) Lisa D. vinealis (Brid.) Zand. var. vinealis D. vinealis (Brid.) Zand. var. flaccidus (Bruch & Schimp. in Schimp.) Zand. Distichium capillaceum (Hedw.) Bruch & Schimp. i n B . S . G . * Ditrichum flexicaule (Schwaegr.) Hamp. D. heteromallum (Hedw.) Britt. D. montanum Leib. Drepanocladus aduncus (Hedw.) Warnst. D. crassicostatus Janssens Dryptodon patens (Hedw.) Brid. Encalypta ciliata Hedw. E. procera Bruch E. vulgaris Hedw. Epipterygium tozeri (Grev.) Lindb. Eucladium verticillatum (Brid.) Bruch & Schimp. in B . S . G . Eurhynchium oreganum (Sull.) Jaeg. E. praelongum (Hedw.) Schimp. in B . S . G . E. pulchellum (Hedw.) Jenn. Fissidens adianthoides Hedw. F. bryoides Hedw. var. bryoides F. bryoides Hedw. var. viridulus (Sw.) Wahlenb. Purs. F. grandifrons Brid. F. limbatus Sull. F. ventricosus Lesq. Fontinalis antipyretica Hedw. var. antipyretica F. antipyretica Hedw. var. gigantea (Sull.) Sull. F. antipyretica Hedw. var. oregonensis Ren. & Card. Funaria hygrometrica Hedw. F. muhlenbergii Turn. 212  Grimmia laevigata (Brid.) Brid. G pulvinata (Hedw.) Sm. G. torquata Hornsch. in Grev. G. trichophylla Grev. Gymnostomum aeruginosum Sm. Hedwigia stellata Hedenas Heterocladium macounii Best. H. procurrens (Mitt.) Jaeg. Homalothecium aeneum (Mitt.) Lawt. H. arenarium (Lesq.) Lawt. H. fulgescens (Mitt, ex C . Mull.) Lawt. H. nuttallii (Wils.) Jaeg. H. pinnatifldum (Sull. & Lesq.) Lawt. Hygrohypnum bestii (Ren. & Bryhn in Ren.) Broth. Hylocomium splendens (Hedw.) Schimp. in B . S . G . Hymenostylium recurvirostre (Hedw.) D i x . Hypnum circinale Hook. H. cupressiforme Hedw. H. dieckii Ren. & Card, in R o l l . H. subimponens Lesq. Isothecium cristatum (Hampe) Robins. / myosuroides Brid. Leptobryum pyriforme (Hedw.) Wils. Leptodictyum riparium (Hedw.) Warnst. Leucolepis acanthoneuron (Schwaegr.) Lindb. Metaneckera menziesii (Hook, in Drumm.) Steere Mnium marginatum (With.) Brid. ex P. Beauv. M. spinulosum Bruch & Schimp. in B . S . G . Neckera douglasii Hook. N. pennata Hedw. Oncophorus wahlenbergii Brid. Orthotrichum afftne Brid. O. anomalum Hedw. O. consimile Mitt. O. hallii Sull. & Lesq. in Sull. O. lyellii Hook. & Tayl. O. pulchellum Brunt, in Winch. & Gateh. O. rupestre Schleich. ex Schwaegr. O. speciosum Nees in Sturm O. striatum Hedw. O. tenellum Bruch ex Brid. Phascum cuspidatum Hedw. Philonotis capillaris Lindb. in Hartm. P. fontana (Hedw.) Brid. Physcomitrium pyriforme (Hedw.) Hampe Plagiomnium insigne (Mitt.) T. K o p . P. medium (Bruch & Schimp. in B.S.G.) T. K o p . P. rostratum (Schrad.) T. K o p . 213  P. venustum (Mitt.) T. K o p . Plagiothecium denticulatum (Hedw.) Schimp. in B . S . G . P. laetum Schimp. in B . S . G . P. piliferum (Sw. ex Hartm.) Schimp. i n B . S . G . Plagiothecium undulatum (Hedw.) Schimp. in B . S . G . Platydictya jungermannioides (Brid.) Crum Platyhypnidium riparioides (Hedw.) D i x . Pleuridium acuminatum Lindb. Pleurozium schreberi (Brid.) Mitt. Pogonatum contortum (Brid.) Lesq. P. urnigerum (Hedw.) P. Beauv. Pohlia cruda (Hedw.) Lindb. P. longibracteata Broth, in R o l l . P. nutans (Hedw.) Lindb. P. sphagnicola (Bruch & Schimp.) Lindb. & Arnell P. wahlenbergii (Web. & Morh) Andrews Polytrichastrum alpinum (Hedw.) G . L . Sm. Polytrichum commune Hedw. P. for mo sum Hedw. P. juniperinum Hedw. P. piliferum Hedw. P. strictum B r i d . Porotrichum bigelovii (Sull.) Kindb. P. vancouveriense (Kindb. i n Mac.) Crum Pseudobraunia californica (Lesq.) Broth. Pseudoscleropodium purum (Hedw.) Fleisch. i n Broth. Pseudotaxiphyllum elegans (Brid.) Iwats. Pterogonium gracile (Hedw.) Sm. Ptychomitrium gardneri Lesq. Racomitrium aciculare (Hedw.) Brid. R. elongatum Ehrh. ex Frisv. R. ericoides (Web. ex Brid.) Brid. R. heterostichum (Hedw.) Brid. R. lanuginosum (Hedw.) Brid. R. lawtonae Irel. R. occidentale (Ren. & Card.) Ren. & Card. R. pacificum Irel. & Spence R. varium (Mitt.) Jaeg. Rhizomnium glabrescens (Kindb.) T. K o p . R. magnifolium (Horik.) T. K o p . Rhytidiadelphus loreus (Hedw.) Warnst. R. squarrosus (Hedw.) Warnst. R. triquetrus (Hedw.) Warnst. Rhytidiopsis robusta (Hook.) Broth. Roellia roellii (Broth, in Roll) Andrews ex Crum * Sanionia uncinata (Hedw.) Loeske Schistidium agassizii Sull. & Lesq. in Sull. S. apocarpum (Hedw.) Bruch & Schimp. in B . S . G . 214  S. maritimum (Turn.) Bruch & Schimp. in B . S . G . S. rivulare (Brid.) Podp. Scleropodium cespitans (C. M u l l . ) L . K o c h S. obtusifolium (Jaeg.) Kindb. in M a c . & Kindb. S. touretii (Brid.) L . K o c h var. touretii Scleropodium touretii (Brid.) L . K o c h var. colpophyllum (Sull.) Lawt. ex Spahgnum capillifolium (Ehrh.) Hedw. S. fuscum (Schimp.) Klinggr. S. henryense Warnst. & magellanicum Brid. S. palustre L . Sphagnum recurvum P. Beauv. S. rubellum Wils. S. squarrosum Crome S. subsecundum Nees i n Sturm Tayloria serrata (Hedw.) Bruch & Schimp. in B . S . G . * Tetraphis pellucida Hedw. Timmia austriaca Hedw. Timmiella crassinervis (Hampe) L . K o c h Tortella fragilis (Hook. & W i l s in Drumm.) Limpr. T. tortuosa (Hedw.) Limpr. Tortula amplexa (Lesq.) Steere T. laevipila (Brid.) Schwaegr. var. laevipila T. laevipila (Brid.) Schwaegr. var. meridionalis (Schimp.) Wijk & Marg. T. latifolia Bruch ex Hartm. T. muralis Hedw. T. papillosa Wils. in Spruce T. papillosissima (Copp.) Broth. T. princeps De Not. T. ruralis (Hedw.) Gaertn. et al. T. subulata Hedw. Trachybryum megaptilum (Sull.) Schof. Trichodon cylindricus (Hedw.) Schimp. Trichostomopsis australasiae (Grev. & Hook.) Robins. Ulota megalospora Vent, i n R o l l . U. obtusiuscula C . M u l l . & Kindb. in M a c . & Kindb. U. phyllantha Brid. Warnstorfia fluitans (Hedw.) Loeske Weissia controversa Hedw. Zygodon viridissimus (Dicks.) Brid. var. rupestris Lindb. ex Hartm.  215  Appendix L Names of taxa that differ from those in Lawton (1971) or are not treated i n that publication are given below. The name on the left is that used in the thesis. Amblstegium serpens var. juratzkanum Andreaea megistospora Bryoerythrophyllum recurvirostre Bryum ambylodon Bryum dichotomum Bryum flaccidum Coscinodon calyptratus Crumia latifolia Didymodon fallax Didymodon rigidulus var. gracilis Drepanocladus crassicostatus Dryptodon patens Fissidens limbatus Hedwigia stellata Hymenostylium recurvirostre Isothecium myosuroides Leptodictyum riparium Leucolepis acanthoneuron Metaneckera menziesii Phytscomitrium pyriforme Platyhpnidium riparioides Pleuridium acuminatum Pohlia sphagnicola Polytrichastrum alpinum Porotrichum bigelovii Porotrichum vancouveriense Pseudotaxiphyllum elegans Racomitrium Racomitrium lawtonae Racomitrium occidentale Racomitrium pacificum Roellia roellii Sanionia uncinata Schistidium agassizii Schistidium apocarpum Schistidium maritimum Schistidium rivulare Sphagnum Trachybryum megaptilum Trichodon cylindricus Ulota obtusiuscula Warnstorfia fluitans Zygodon viridissimus var. rupestris  Amblystegium juratzkanum not treated in Lawton, see Murray (1987) Didymodon recurvirostris Bryum stenotrichum Bryum bicolor Bryum capillare var. flaccidum Grimmia calyptrata Scopelophila latifolia Barbula fallax Barbula acuta not treated in Lawton, see Janssens (1983a) Rhacomitrium patens Fissidens bryoides Hedwigia ciliata Gymnostomum recurvirostre Isothecium stoloniferum Amblystegium riparium Leucolepis menziesii Neckera menziesii Physcomitrium kellermanii & P. megalocarpum Eurhynchium riparioides Pleuridium subulatum not treated in Lawton, see Shaw (1982) Polytrichum alpinum Porothamnium bigelovii Bestia vancouveriensis Isopterygium elegans Rhacomitrium not treated in Lawton, see Ireland (1970) or Frisvoll (1988) Rhacomitrium heterostichum var. occidentale not treated in Lawton, see Ireland & Spence (1987) or Frisvoll (1988) Bryum sandbergii Drepanocladus uncinatus Grimmia alpicola Grimmia apocarpa Grimmia maritima Grimmia alpicola var. rivularis not treated in Lawton, see Crum (1984) Homalothecium megaptilum Ditrichum cylindricum Ulota crispa var. alaskana Drepanocladus fluitans Zygodon vulgaris  216  Appendix M Species that could be expected to be in the San Juan Islands but were not found. Barbula convoluta Hedw. Bartramia stricta Brid. Calliergon stramineum (Brid.) Kindb. Campylopus fragilis (Brid.) Bruch & Schimp. in B . S . G . C. subulatus Schimp in Rabenh. Dicranella rufescens (With.) Schimp. D. cerviculata (Hedw.) Schimp. Ditrichum ambiguum Best. Encalypta rhaptocarpa Schwaerg. Entosthodon fascicularis (Hedw.) C . M u l l . Hygrohypnum ochraceum (Turn, ex Wils.) Loeske Oligotrichum aligerum Mitt. Plagiothecium cavifolium (Brid.) Iwats. Pohlia proligera (Kindb. ex Breidl.) Lindb. ex Arnell Pottia truncata (Hedw.) Furnr. ex B . S . G . Racomitrium obesum Frisv. Sphagnum mendocinum Sull. & Lesq. in Sull.  217  Appendix N County reference map and distribution maps for all o f the species.  218  220  Andreaea megistospora  Andreaea rupestris 221  222  224  Brachythecium rivulare  Brachythecium velutinum var. velutinum  Brachythecium velutinum var. venustum 225  Bryoerythrophyllum recurvirostre  Bryum caespiticium  Bryum canariense 226  227  Bryum lisae var. cuspidatum  Bryum miniatum  Bryum pallens  Bryum pallescens 228  Calliergon giganteum  Calliergonella cuspidata 229  Dendroalsia abietina  Desmatodon obtusifolius 232  233  234  236  Ditrichum montanum  Drepanocladus aduncus  Drepanocladus crassicostatus  Dryptodon patens 237  Encalypta ciliata  Encalypta procera  238  239  Fissidens bryoides var. viridulus  Fissidens grandifrons  240  Fontinalis antipyretica var. antipyretica 241  Fontinalis antipyretica var. gigantea  242  Grimmia trichophylla  Gymnostomum aeruginosum  243  Heterocladium procurrens  Homalothecium aeneum 244  245  246  247  Neckera douglasii  Neckera pennata  249  Orthotrichum pule helium  Orthotrichum rupestre  251  252  Philonotis capillaris  Philonotis fontana  Physcomitrium pyriforme  Plagiomnium insigne  253  254  Plagiothecium laetum  Plagiothecium piliferum  255  256  Pohlia longibracteata  Pohlia nutans 257  258  Polytrichum piliferum  Polytrichum strictum 259  Pseudobraunia californica  Pseudoscleropodium purum 260  Ptychomitrium gardneri  Racomitrium aciculare 261  Racomitrium elongatum  Racomitrium ericoides  Racomitrium heterostichum  Racomitrium lanuginosum 262  Rhytidiadelphus lore us  Rhytidiadelphus squarrosus 264  265  Schistidium apocarpum  Schistidium maritimum  Schistidium rivulare  Scleropodium cespitans  266  Scleropodium obtusifolium  Scleropodium touretii var. touretii  Scleropodium touretii var. colpophyllum 267  Sphagnum capillifolium  268  269  271  Tortula papillosa  Tortula papillosissima 272  274  Ulota phyllantha  Weissia controversa  Warnstorfia fluitans  Zygodon viridissimus var. rupestris  . 275  

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