Inventory of the Mosses, Liverworts, Hornworts, and …Inventory of the Mosses, Liverworts,...
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Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
U.S. Department of the InteriorU.S. Geological Survey
Scientific Investigations Report 2005-5240
Prepared in cooperation with Olympic National Park
Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
By Martin Hutten Andrea Woodward
Karen Hutten
U.S. Department of the InteriorU.S. Geological Survey
Scientific Investigations Report 2005–5240
U.S. Department of the InteriorGale A. Norton, Secretary
U.S. Geological SurveyP. Patrick Leahy, Acting Director
U.S. Geological Survey, Reston, Virginia: 2005
For sale by U.S. Geological Survey, Information Services Box 25286, Denver Federal Center Denver, CO 80225
For more information about the USGS and its products: Telephone: 1–888–ASK–USGS World Wide Web: http://www.usgs.gov/
Any use of trade, product, or firm names in this publication is for descriptive purposes only and does not imply endorsement by the U.S. Government.
Suggested citation:
Hutten, Martin, Woodward, Andrea, and Hutten, Karen, 2005, Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List. U.S. Geological Survey, Scientific Investigations Report 2005-5240, 78 p.
Key words:
1. US Geological Survey. 2. National Park Service. 3. Olympic National Park. 4. Non-vascular Plants. 5. Inventory. 6. Environmental sampling. 7. Rare Plants. 8. Coniferous forests. 9. Species list. 10. Cryptogams. 11. Bryophytes. 12. Mosses. 13. Liverworts. 14. Hornworts.
iiiAcknowledgments
Acknowledgments.
This project has been funded jointly by the U.S. Geological Survey, Canon USA, Inc., through the National Park Foundation, and Olympic National Park. Volunteers Jeanette and Paul Stehr-Green, Kelli Weed, Anita Woodnut, and Lisa McMillan dedicated hundreds of hours to this project. Steven Hahn, Lisa Nesbitt, Krista Rome, James Walton, and Kelli Weed helped with field and office work. Ed Schreiner was involved in early discussions of the sampling plan. Bruce McCune and others helped with identification of difficult lichens.Gay Hunter helped with the herbarium collections and Paul Gleeson provided a workspace with quality microscopes, close to the collections. Dan Norris provided much enthusiasm that proved contagious. The authors were further encouraged by Dr. Judith Harpel, Dr. Wilfred Schofield, and Dr. Tør Tønsberg. Tør also added more than 50 lichens to the Olympic Peninsula lichen list.
iv Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Units of Measure for Conversions1 meter = 39.37 inches1 kilometer = 0.62 miles1 square centimeter (cm2) = 0.15 square inches (in2)1 square kilometer (km2)= 0.39 square miles (mi2)
vContents
ContentsAcknowledgements ..................................................................................................................................... iiiUnits of Measure for Conversions ............................................................................................................ ivAbstract .......................................................................................................................................................... 1Introduction ................................................................................................................................................... 1
Geographic and Biologic Context for the Inventory ...................................................................... 2Signifi cance of Cryptogams for ONP ................................................................................................ 2Project Description .............................................................................................................................. 3
Methods ......................................................................................................................................................... 3Field Methods ....................................................................................................................................... 3
Modifi ed FHM Method ...................................................................................................... 3Intuitively Controlled Method .......................................................................................... 4
Processing of Specimens ................................................................................................................... 5Equipment and Chemicals ............................................................................................... 5Processing Method ........................................................................................................... 5Identifi cation Literature ................................................................................................... 5Names and Titles of Experts Who Were Consulted .................................................... 7
Curatorial Methods .............................................................................................................................. 7Archival-Quality Voucher Envelopes .............................................................................. 7Database ............................................................................................................................. 8Quality Assurance/Quality Control for Data Entry ........................................................ 8Storage ................................................................................................................................ 8
Results ............................................................................................................................................................ 8Species List ........................................................................................................................................... 8Other products .................................................................................................................................. 53
Field Guide ........................................................................................................................ 53Illegal Commercial Moss Harvests ............................................................................... 53Fact Sheet ......................................................................................................................... 53Websites ............................................................................................................................ 53GIS Layer ........................................................................................................................... 53Metadata ........................................................................................................................... 53Collaboration with Other Agencies ............................................................................... 53
Discussion .................................................................................................................................................... 55Cryptogam Diversity of the Olympic Peninsula ............................................................................ 55Rare Species on the Olympic Peninsula ........................................................................................ 55Endemic Species .............................................................................................................................. 56
Regional Context for Rare Species .............................................................................. 56Threats ............................................................................................................................... 57
Further Work in Olympic National Park ......................................................................................... 59Needs for Existing Bryophyte and Lichen Data ......................................................... 59Needed Field Work ......................................................................................................... 61
Conclusions ................................................................................................................................................. 67References ................................................................................................................................................... 67Appendix A ................................................................................................................................................... 71
vi Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
FiguresFigure 1. Collection localities on the Olympic Peninsula ................................................................... 4Figure 2. Example of the information on a typical voucher ............................................................... 7Figure 3. Part of the lichen collection at the Olympic National Park curatorial facility ............... 9Figure 4. Diplophyllum plicatum ........................................................................................................... 54Figure 5. Herbertus aduncus ................................................................................................................ 54Figure 6. Bundophoron melanocarpum .............................................................................................. 54Figure 7. Pseudocyphellaria mallota ................................................................................................... 54Figure 8. Splachnum ampullaceum. .................................................................................................... 54Figure 9. Tayloria serrata ...................................................................................................................... 58Figure 10. Blepharostoma arachnoideum ............................................................................................ 58Figure 11. Pseudocyphellaria rainierensis ........................................................................................... 58Figure 12. Splachnum sphaericum ........................................................................................................ 58Figure 13. Schistostega pennata ........................................................................................................... 58Figure 14. Tetraphis geniculata .............................................................................................................. 58Figure 15. Bryoria cf. fremontii ............................................................................................................... 60Figure 16. Usnea sphaecelata ................................................................................................................ 60Figure 17. Umbilicaria rigida ................................................................................................................... 60Figure 18. Orthodontium lineare ............................................................................................................ 60Figure 19. Campylopus introflexus ........................................................................................................ 60Figure 20. Ahtiana sphaerosporella. ..................................................................................................... 60Figure 21. Gymnomitrion obtusum ......................................................................................................... 64Figure 22. Iwatsukiella leucotricha ....................................................................................................... 64Figure 23. Racomitrium aquaticum ........................................................................................................ 64Figure 24. Rhytidium rugosum. ............................................................................................................... 64Figure 25. Crumia latifolia ....................................................................................................................... 64Figure 26. Fissidens grandifrons ............................................................................................................ 66Figure 27. Hydrothyria venosa ............................................................................................................... 66Figure 28. Pilophorus nigricaulis ........................................................................................................... 66Figure 29. Discelium nudum ................................................................................................................... 66Figure 30. cf. Tetraplodon mniodes ....................................................................................................... 66Figure 31. Athalamia hyalina. ................................................................................................................ 66
TablesTable 1. Species list of mosses, liverworts, hornworts and lichens and associated
vegetation types and substrate in Olympic National Park, Washington. ................. 10Appendix A Rare bryophytes and lichens known from Olympic Peninsula ................................... 71
1Olympic Botanists, 441 Hudson Road, Port Angeles, WA 98363, [email protected]
2USGS Forest & Rangeland Ecosystem Science Center Olympic Field Station, 600 E. Park Avenue, Port Angeles, WA 98362
Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
By Martin Hutten1 Andrea Woodward2
Karen Hutten1
AbstractThe identification of non-vascular cryptogam species
(lichens, mosses, liverworts, and hornworts) is especially chal-lenging because of their small size, their often microscopic or chemical distinguishing features, and their enormous diversity. Consequently, they are a poorly known component of Olympic National Park, despite their ecological and aesthetic impor-tance. This project is the first attempt at a systematic, compre-hensive survey of non-vascular cryptogams in the Park and presents the current species list with descriptions of the sub-strate and vascular vegetation type where they were observed. The authors strove to collect from as many park environ-ments as feasible, and distributed collections along important environmental gradients in different regions of the park using vascular vegetation as an environmental indicator. They also collected opportunistically when interesting habitats or microhabitats were encountered. Finally, the authors updated the nomenclature in the Park’s previous collection of non-vascular plants. This study identified approximately 13,200 bryophyte and lichen species, adding approximately 425 new species to the Olympic National Park Herbarium. These data, combined with select literature reports and personal data from Martin and Karen Hutten, added more than 350 species to the previously documented Olympic Peninsula lichen and bryo-phyte list. The authors discuss the list in a local, regional, and global context of rarity, as well as cryptogam conservation and further work needed in Olympic National Park. The improved inventory of Olympic National Park cryptogams represented by this project enables Olympic National Park to protect popu-lations of rare and sensitive species, assess the damage caused by illegal harvest, and contribute information to the Bureau of Land Management and U.S. Forest Service Sensitive Species Programs.
IntroductionThe identification of cryptogam species (lichens, mosses,
liverworts, and hornworts) is especially challenging because of their small size, their often microscopic or chemical distin-guishing features, and their enormous diversity. Consequently, they are a poorly known component of Olympic National Park (ONP), despite their ecological and aesthetic importance. Although experts have created partial species lists for ONP (Sharpe, 1956; Harthill, 1964; Thomson, 1969; Kunze, 1980; Hong and others, 1989) based on surveys of limited areas, a systematic, comprehensive survey has not been conducted. The improved inventory of ONP cryptogams represented by this project enables ONP to protect populations of rare and sensitive species, assess the damage caused by illegal harvest, and contribute information to the Bureau of Land Management (BLM) and U.S. Forest Service (USFS) Sensitive Species Programs.
This document presents the current species list of mosses, liverworts, hornworts, and lichens of ONP, along with descrip-tions of substrate and vascular vegetation type where they were observed. The authors discuss the list in a local, regional, and global context of rarity, as well as address some general topics regarding cryptogam conservation and further work needed in ONP. The authors do not believe it to be a complete list, nor do they analyze or summarize species distribution or community structure.
Although ONP comprises slightly less than 10 percent of the Olympic Peninsula (OP), it protects the bulk of the unmanaged ecosystems and thereby represents the flora that may once have been present on the entire OP. Because some collections and some of the records in the Park’s herbarium come from outside of the Park, conclusions and generaliza-tions are sometimes applied only to ONP and sometimes to the entire OP.
Geographic and Biologic Context for the Inventory
ONP is the centerpiece of the OP, which is a 13,800-km2 landmass in the extreme northwest corner of the contermi-nous United States (fig. 1). The Peninsula resembles an island because it is surrounded on three sides by water and on the fourth by a deep river valley: the Pacific Ocean to the west, the Strait of Juan de Fuca to the north, Hood Canal to the east, and the Chehalis River Valley to the south. The Olym-pic Mountains (Olympics) rise from sea level to culminate at Mount Olympus near the center of the Peninsula at 2,430 m. Geologic uplift, heavy precipitation, and a dynamic glacial history have created a radial pattern of 11 major river valleys centered on the mountains. ONP covers 3,700 km2 in two units: 3,530 km2 in the central mountainous core and a nar-row 170-km2 strip extending 84 km along the coast (Olympic National Park, 1999).
Mountainous areas in general are characterized by steep moisture and temperature gradients, resulting in substantially different environments over short distances. The moisture gradient is especially steep in the Olympic Mountains, reflect-ing the wet maritime climate, and results in exceptionally high levels of precipitation along the western slope. The substan-tial climatic and elevation gradients of the Peninsula create a diversity of habitats within the park. Environmental regimes range from mild, maritime conditions on the coast to harsh, alpine areas at high elevations to dry, near-continental climate in the northeast. Consequently, cold-stressed alpine vegetation exists within 15 km of intertidal communities and even closer to lush temperate rainforest with some of the world’s largest trees (Buckingham and others, 1995).
The geologic and glacial histories of the Peninsula and western Washington provide a diversity of parent materials for soil formation. The ocean floor contributed sedimentary and marine-deposited basaltic bedrock. The continental glaciers deposited a variety of soil material including granitic rocks from the Cascade Range along the east and north sides of the Peninsula. Mass wasting and glaciers have mixed, washed, and eroded all three materials, creating a complex of montane and riverine soil materials (Tabor, 1987). Olympic soils are considered to be young and, in general, are relatively infer-tile, except in the lower Dungeness River Valley. Local soil characteristics (for example, soil moisture, sub-surface flow, soil temperature, and chemical properties) vary greatly, being influenced by the parent material, climate, and biotic com-munities of the area. Common soil orders include spodosols, inceptisols, entisols, histosols, and andisols (Henderson and others, 1989).
Vegetation reflects the diverse physical environment. West-side lowland forests are in the Sitka Spruce (Picea sitchensis) Zone, including the temperate coniferous rainfor-est for which ONP is famous. Here, massive Sitka spruce trees grow to 90 m, and deciduous bigleaf maples (Acer macrophyl-lum) are laden with epiphytes. Lowland and mid-elevation forests on the drier east side and mid-elevation forests on the
west side are in the Western Hemlock (Tsuga heterophylla) Zone. This is the most widespread zone and it is dominated by Douglas-fir (Pseudotsuga menziesii) and western hemlock, with red cedar (Thuja plicata) as a fairly common constituent. Montane forests are in the Pacific Silver Fir (Abies amabi-lis) Zone on the cool, moist slopes of much of the Peninsula, whereas Douglas-fir inhabits south-facing montane slopes in the northeast. Subalpine areas are a matrix of tree islands and meadows at high elevation. Subalpine areas with snow packs deeper than 3 m are in the Mountain Hemlock Zone (Tsuga mertensiana) and include mountain hemlock, subalpine fir (Abies lasiocarpa), and sometimes Pacific silver fir. The Subalpine Fir Zone occurs in areas with snowpacks less than 3 m deep and may also include lodgepole pine (Pinus contorta) or whitebark pine (P. albicaulis). Treeline occurs at 1,615 m in wetter areas and 1,890 m in drier zones where trees finally give way to alpine meadows (Henderson and others, 1989).
The glacial history, geographic isolation, and steep cli-matic gradients have important consequences for the biogeog-raphy of the area. First, the Peninsula was never completely covered by ice during the last glaciation at least, when a complex of ridges and mountains were above ice. Moreover, when glacial ice was thick, sea level was lower, exposing con-siderable new lands along the coast for a long period of time (Booth, 1987; Tabor, 1987). The role of the OP as a glacial refugium is conjecture, but the theory is well supported by its biogeography (Buckingham and others, 1995). The OP is home to a surprising number of endemic and disjunct vascular species whose distribution patterns are consistent with the refugium theory. Not enough is known about the distribution of non-vascular species to reach conclusions, but the disjunct distribution patterns of several species support the theory (for example, Rhytidium rugosum and Vulpicida tilesii are both arctic Tertiary relics now disjunct from their main distributions).
Significance of Cryptogams for ONP
Non-vascular cryptogams (mosses, liverworts, and lichens) make up a significant portion of the biomass and annual production in temperate rainforests (Vitt and others, 1988), and are known to play important ecological roles in hydrologic (Norris, 1990; Boucher and Nash, 1990) and nutrient cycling (Nadkarni, 1985). They are also known to be sensitive to precipitation chemistry and air quality (Hawk-sworth and Hill, 1984). Consequently, non-vascular cryp-togams are crucial components of the long-term ecological monitoring program currently under development in ONP. Cryptogams require management attention because they are harvested illegally for sale to the floral industry, and because some rare species may require special protection.
Additionally, mosses, liverworts, and lichens were included in the list of Survey and Manage species identified in the Record of Decision (U.S. Department of Agriculture and U.S. Department of the Interior, 1994) for the Northwest
2 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Forest Plan, to which the National Park Service (NPS) is a party. The Northwest Forest Plan was adopted by the BLM and USFS in 1994 to guide forest management in the range of the northern spotted owl after the owl’s designation as endangered under the Endangered Species Act. Part of the plan required the Forest Service to survey for certain species (in other words, Survey and Manage Species) before allow-ing ground-disturbing activities. The Survey and Manage List included a large number of bryophytes and lichens, among other taxa, and was based on minimal knowledge of the dis-tribution and abundance of these species. ONP can fulfill its obligation to aid regionally based management of bryophytes and lichens mandated by the USFS and BLM Sensitive Spe-cies Programs by providing information about the status of bryophyte and lichen populations within its borders.
Project Description
There have been few previous attempts to systematically describe the biodiversity of cryptogams in ONP. Collectors have explored specific areas or places within short walking distance of parking lots. Consequently, the authors strove to expand collections to as many park environments as feasible and distribute them along important environmental gradients in different regions of the park using vascular vegetation as an environmental indicator. They also collected opportunistically when interesting habitats or microhabitats were encountered. Finally, the authors updated the nomenclature in the Park’s previous collection of non-vascular plants. Because of the size of this collection (approximately 5,000 specimens), only obvi-ous misidentifications were corrected. The outcome of this project is a more comprehensive survey and more complete species list than previously available, documentation of cryp-togam communities, and improvement of the Park’s herbarium collection.
Methods
Field Methods
Two types of field methods were employed over three field seasons during this project. In 1999 data were collected intuitively, by searching for specific types of microhabitats. Select diversity hot spots were visited to collect and pho-tograph rare cryptogams. A more systematic inventory of selected major vegetation types was begun in 2000 of Sitka Spruce, Douglas-fir, dry Western Hemlock, moist Western Hemlock, dry Silver Fir, moist Silver Fir, Mountain Hem-lock, and Subalpine Fir Zones and alpine parkland throughout the Park. Data were collected according to a modified U.S. Department of Agriculture (USDA) Forest Service Forest Health Monitoring (FHM) protocol (U.S. Forest Service, 1999, described below). Incidental collections and observa-tions using the intuitive approach were also made in 2000. Field methods in 2001 were the same as during 2000, but data were collected in only a few plots.
• Modified FHM MethodSurvey sites were selected to include major environmen-
tal gradients in ONP and to achieve a distributed geographic sample by using vascular vegetation to indicate environmen-tal conditions. Because there is no map of ONP vegetation at the desired plant-community scale, the Potential Natural Vegetation Map (Henderson and Peters, 2000) and the Park’s geographic information system (GIS) were used to stratify ONP by selected major vegetation types. Foot trails that pass through areas of the park having the highest diversity of major vegetation types were selected as sampling routes. The authors planned to complete three plots in each selected vegetation type during the field season. Plots were selected by locating five random points (changed to 10 later in field season) within 200 m of the trail in the section with the target vegetation types. Areas having slopes of more than 35o were rejected at this stage for safety reasons. The Universal Transverse Mercator (UTM) coordinates of the random points were plot-ted on maps along with an associated random priority ranking for each. A two-person field crew used a global positioning system (GPS) unit to locate collection points. Data were col-lected at the highest priority UTM point if actual vegetation type matched projected vegetation type and if the collection site was safe to sample. If the vegetation was incorrect or the site unsafe, the crew continued to the next priority site until the target number of plots was completed. The final selection of plots produced a well-distributed and diverse sample of the Park (fig. 1).
Methods 3
• Intuitively Controlled MethodMany bryophytes and lichens are highly microhabitat-
specific. Whereas microhabitats are easily targeted when encountered in the field, they are often missed in a sampling method based on more general macrohabitats. Consequently, an intuitively controlled sampling method is more effective in locating certain rare microhabitat- and substrate-specific species. For example, some species of cyano-lichens are found nearly exclusively on semi-exposed conifer twigs in perennially cool and moist microhabitats. Typically, they are further restricted to older forest, in areas where the common cyano-lichens are particularly abundant. When such places were encountered en route between randomly selected plots in the back country, they were checked for the presence of such specialized species if time allowed.
Data were collected using the lichen protocol employed by USFS for the FHM plots. It is a time-constrained search method for epiphytic macrolichens in 36.6-m radius plots for up to 2 hours (U.S. Forest Service, 1999) or until no new spe-cies have been found in 10 minutes. The authors modified this approach in several ways.
1. Instead of limiting the sample to epiphytic lichens, field observers searched for all macrolichens, includ-ing terrestrial lichens; crustose lichens were collected only accidentally or randomly. Although this modified method leads to a somewhat larger sampled commu-nity than the FHM method, the diversity of terrestrial lichens is not high in most heavily forested vegetation types. Within parkland plots, essentially all species are terrestrial, so parklands were a significant departure
Figure 1. Collection localities on the Olympic Peninsula. Collection localities marked with orange dots are associated with extensive Forest Health Monitoring (FHM) type community data, generated in 2000 and 2001. Localities marked with yellow have collections (one to many) generated from non-FHM search protocols. Most collections outside Olympic National Park are personal data collected by M. Hutten.
4 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
6. It would have been impossible to pack out voucher specimens for all species on longer trips. Therefore, the authors allowed observational data to be collected for species that can be identified with confidence in the field. Nevertheless, if an easily identified species was interesting for any reason (for example, rare, eleva-tion minimum or maximum, or a first collection in a geographic area in ONP), a voucher specimen was collected.
Processing of Specimens
Upon returning from the field, specimens were air dried. Some liverwort genera with ephemeral identification charac-ters (for instance, oil bodies) were placed in a refrigerator, but refrigeration capacity was quickly exceeded. As a result, many liverworts had to be identified without oil-body characteristics. Specimen data were entered into a database. Dried vouchers were filed alphabetically in their respective taxonomic group (lichens, mosses, and liverworts/hornworts). Specimens were placed in a temporary holding area in the ONP herbarium, in bryophyte boxes with labeled tabs marking genera and species.
Identification Methods
• Equipment and Chemicals Standard tools and methods were used to identify bryo-
phytes (Schofield, 1985) and lichens (McCune and Geiser, 1997; White & James, 1988). Identifications were made by M. and K. Hutten, with some specimens sent to taxonomic experts for verification.
• Processing MethodCryptogam specimens were grouped by genus and identi-
fied one genus at a time. This allowed the determinations to proceed efficiently, building familiarity with the diversity of characteristics within each genus and the available generic key. Importantly, this processing method increased the deter-mination accuracy, especially for difficult genera. Associated species frequently were identified as well. Many of the small-est lichens and liverworts overlooked in the field are often associated with larger species in voucher specimens. In some plots the associated species contributed 30 percent to the over-all plot list. When an interesting associated species was found during determinations, the voucher would be refiled under the generic name of the associate, and identified later with others of that genus. Identifications within previously completed genera were made without delay.
• Identification Literature Initially, “Moss Flora of the Pacific Northwest,” by Elva
Lawton (1971), was the basis for specimen identification. All
from the conditions for which the FHM method was designed. This new environment did not create any problems.
2. Because the main goal of this project was to increase understanding of non-vascular plant occurrence in ONP, the 2-hour time constraint built into the FHM method was not fully appropriate. The authors sus-pected that 2 hours would not be sufficient to sample the bryophyte community completely. Therefore, if the observer felt it necessary, sampling could be con-tinued beyond 2 hours (this rarely happened). For each specimen the 15-minute time interval of collection was recorded, enabling these data to be compared with FHM data.
3. With the relatively high diversity of macrolichens in ONP’s mature forests, the authors suspected that sampling bryophytes and macrolichens simultaneously would be too demanding of one observer. Typically, two people would work on a single plot simultane-ously, one taking lichen data, the other bryophyte data. At other times it was deemed most efficient for the crew to split up so that each observer would complete an entire plot alone (4 hours).
4. For lichens, the FHM abundance ratings are based on the frequency of individual thalli.
1 = rare (less than four individuals)
2 = occasional (four to 10 individuals
3 = common (more than 10 individuals)
4 = more than half of the branches and trunks seen have this species
Bryophyte data (mosses, liverworts, and hornworts) were collected analogous to lichen data. To apply the lichen abundance rating to bryophytes, however, requires a slightly different concept of ‘individual’. For bryophyte species that grow in large mats, the authors defined an individual as a patch of approximately 10 cm2. This follows a concept used by Abbey Rosso (2000) in western Oregon.
5. It was not expected that the FHM method would be effective at discovering rare species that occur only in specialized microhabitats. Consequently, the authors planned to use an intuitively controlled method to search for species not found in the FHM plot, but in the same target vegetation and nearby the structured plots. The authors called these extra-plot searches. Conducting intensive searches in areas with the high-est potential for locating a target species is a method employed by the USFS and BLM (U.S. Department of the Interior, 1998). Because overall trip goals tended to be ambitious, extra-plot searches were not always done thoroughly, and sometimes were omitted.
Methods 5
other literature was acquired as the project continued and the need for better keys arose.
MossesSubclass Sphagnidae: Crum (1984)
In 2004 a new key became available (Zander, 2004), which includes at least two species not reported in Crum that are found on the OP: Sphagnum pacificum (Flatberg, 1989) and Sphagnum alaskense (Andrus & Jenssens, 2003). Using the old keys, S. pacificum would key to S. recurvum s.l. During this project we often determined S. recurvum var. brevifolium. This is a problematic taxon for western North America (W. B. Schofield, written commun., 2004). For these reasons, all collections in the S. recurvum group need to be redetermined. S. alaskense was recently reported from ONP, but it is not likely the authors overlooked this species in their collections. S. cuspi-datum, determined twice in this project, is misidenti-fied (W. B. Schofield, written commun., 2004).
Subclass Andreaeidae: Lawton (1971); Murray (1986, 1988) The authors initially used Lawton (1971) but later
switched to Murray (1986, 1988). At some time in the future, the Andreaea identifications should be verified, identifying any newer species not included in Lawton (1971) that may have been overlooked. This probably resulted in the misdetermination of A. megistospora as A. rothii (W.B. Schofield, written commun., 2004). A subset of the substantial Harthill collections, predating this project and kept at ONP, should also be redetermined.
Subclass Bryidae: Lawton (1971) For the following select genera in this large subclass
we used more recent keys as listed after each genera: Pohlia (Shaw, 1982), and Didymodon (Zander, 1999). The complex genus Racomitrium was started using Lawton (1971), but later the authors switched to Frisvoll (1983, 1988), in which several species aggregates are differentiated. As a result, a portion of the Racomitrium collections should be re-exam-ined at some time in the future, as well as a subset of the substantial Harthill collections predating this project that are kept at ONP. The Frisvoll volumes do not cover all species in the genus Racomitrium, so we used Lawton (1971) for the remainder of the species. At present Norris and Shevock (2004a) should be considered (see also Bednarek-Ochyra, 2000). Dicranum species were verified using Lawton (1971), and for Washington this key is probably adequate. Rare species were verified using a recently published key (Zander 2004), which is more up to date. The many ONP Dicranum bonjeanii collec-tions in the ONP Herbarium submitted by Harthill
and others are probably Dicranum howellii and D. scoparium. These should be verified using the new Dicranum key in Zander (2004). Some species con-cepts/groups have been revised fairly recently and our ONP specimens should therefore be re-examined (in part) at some point in the future, including Hedwigia ciliata; ours is now called H. stellata (Buck & Norris, 1996).
LiverwortsA complete key to liverworts and hornwort (hepatics)
genera recently became available (Schofield, 2002). Unfortu-nately, this work does not contain keys to the species level. At present there is no single work that can be used to identify all hepatic species that occur in ONP. For keys and descriptions of most species found at ONP, one of the best single resources is Judith Godfrey’s thesis (1977). The authors used this work extensively, along side updated versions of the Christy and Wagner (1996) keys, the Schuster volumes (1966-1992), and versions of hepatic keys by Dan Norris (unpublished). Rather late in the identification process the authors acquired “The Liverwort Flora of the British Isles” (Paton, 1999), which helped resolve many problems in segregating taxa using the keys in the resources listed above. The “Illustrated Flora of Nordic Liverworts and Hornworts” (Damsholt 2002) would also be useful for some genera, but unfortunately was unavail-able. None of the above resources cover all species found in ONP, consequently a myriad of additional literature is needed to identify some of the lesser-known species. Some of this literature was not available to this project.
LichensFortunately, the literature relevant to macrolichens is
much more consolidated than is the bryophyte literature. The keys in McCune and Geiser (1997) cover all but a few macro lichens. In general, the omitted lichens are more northern species that can be found in Goward (1999) and Goward and others (1994). For Leptogium, Pseudocyphellaria, Usnea, and Xanthoria (McCune, 2000), the authors used updated keys available from Bruce McCune (2000).’s website: (http:// oregonstate.edu/~mccuneb/getkeys.htm). Some species concepts/groups have been revised fairly recently and ONP specimens should therefore be re-examined (in part) at some point in the future, including Pannaria/Fuscopannaria, Menegazzia, Physconia (P. fallax is a species now known from the ONP, but was not in the keys used by the authors for the ONP material), Pyrrhospora, and so forth. Most crustose lichens were omitted in this project, but can be keyed to genus using McCune (2002). Many crustose lichens can be keyed to species using Noble (1982) and Purvis and others (1992).
6 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
• Names and Titles of Experts Who Were Consulted The following people have verified specimens during this project. Bruce McCune provided the majority of lichen verification.
1. Dr. Rick Dewey, USFS Botanist: Riccia, Tritomaria
2. Chiska Derr, USFS lichen Taxa Expert: Survey and Manage lichens
3. Dr. Linda Geiser, USFS Ecologist and Air Quality Specialist: Bryoria capillaris
4. Dr. Katie Glew, University of Washington Herbarium (WTU): Alectoria samentosa subsp. vexillifera
5. Dr. Judith Harpel, Regional Interagency Bryologist: Survey and Manage bryophytes
6. Sarah Jovan, Ph.D. candidate, student under McCune: Physconia fallax
7. Dr. Bruce McCune, Professor, Oregon State University (OSU): lichens
8. Dr. Wilfred Schofield, Professor, University of British Columbia (UBC): bryophytes
Curatorial Methods
• Archival-Quality Voucher EnvelopesArchival-quality voucher envelopes were produced show-
ing standard collection information for each species (fig. 2). A map of the plot locality is also printed on the inside of each voucher envelope. Both the label and the map are printed on the laserprinter in grayscale on acid-free paper.
Figure 2. Example of the information on a typical voucher envelope generated during this project.
Methods 7
• DatabaseA back-end database retains and provides for the integrity of all raw data. See metadata for a detailed description of back-end database structure (http://mercury.ornl.gov/nbii/full.jsp?index=0&recidx=0&pageMax=1, November 2005). A front-end database was developed by M. Hutten to create data summaries and additional utilities.
• Quality Assurance/Quality Control for Data EntryVarious database tools were employed to ensure accurate
and complete data entry. For example, certain mimimum col-lection-site data are required before the database will connect voucher data to that site. Similarly, certain collector data must be entered before a collector can be connected to a voucher. A voucher record that lacks these and other critical fields cannot be saved until problems are corrected. The user will be prompted to add the missing data. In several fields, such as the abundance rating, the entry is limited to pre-defined valid entries. The data entry in such fields is usually via drop-down lists from which the applicable entry is selected, expediting the data-entry process, and reducing data-entry errors. These drop-down lists can be embedded directly into the defini-tion of the data table or be associated with specific data-entry forms. More extensive, dynamic drop-down lists are easier to handle as a (lookup) data table. When the specimens had been identified and the data entered, new voucher labels were printed on acid free paper. If key fields are lacking or out of bounds (for example, elevations above 3,000 m), the database does not allow the label to be printed. Printed label data were compared and verified with the field label and associated iden-tification notes before transferring the specimens in the new archival-quality envelope.
• StorageSpecimens are stored at Olympic National Park, 600 Park
Avenue, Port Angeles, WA 98362-6798, USA (fig. 3).
ResultsBryophyte and lichen data were collected from 65 modi-
fied FHM-style plots throughout ONP (fig. 1) as the main focus of the project. Less formal data collection occurred opportunistically and intuitively from other localities in ONP during the project. The bulk of the opportunistic collections were made near trails and roads while traveling to plot loca-tions. The scope of these collection efforts ranged from a single interesting species found along a trail, to substantial col-lections from an area of interest. Many of the ‘other protocol plots’ indicated on figure 1 are localities that were visited by M. and K. Hutten during trips not related to this project. These data are included in the species list and map because they provide additional documentation of the OP bryophyte and lichen flora.
Species List
During the course of this study, some 13,200 bryophyte and lichen species were identified (table 1), adding approxi-mately 425 new species to the ONP Herbarium. These data, combined with select literature reports and personal data from M. and K. Hutten, added more than 350 species to the already extensive OP lichen and bryophyte list compiled from litera-ture by Fred Rhoades (written comm., 1997). A few species were removed from Rhoades’ list because the reports were not based on actual collections, or they were based on misdeter-minations (for example, all reports of Scouleria marginata; J. Harpel, oral commun., 2002). Verification of literature reports requires finding and verifying the determinations of voucher specimens that the literature reports were based on. This was beyond the scope of this project, and it is therefore conceivable that some species listed on the Rhoades’ list are in error. Species such as Rhizomnium appalachianum, Scapania nemorosa, and Telaranea nematodes were removed, as they are known exclusively from eastern North America. Cur-rently, more than 1,280 non-vascular plant species, subspecies, and varieties have been documented from the OP (table 1). Some reports still may contain errors, but the authors made no attempt to verify any specimens that were not available at ONP. Only a number of the approximately 5,000 non-vascu-lar plants in the ONP collections predating this project were verified. Checking previous collections was not a goal of this project, and typically records were examined only when errors were suspected. Therefore, many determinations in the ONP collection still need verification. Moreover, some of our deter-minations need further work for several reasons. Some groups of non-vascular plants were challenging to identify, in part due to the lack of complete and accurate keys for all species in the Pacific Northwest (PNW). Ecological and distributional infor-mation for many species, especially bryophytes, is still very incomplete. Plot work often yields many imperfect specimens, which may be small, lack needed structures (for example, sexual), or specimens were not examined when sufficiently fresh (for instance, some liverworts have ephemeral charac-ters), making identification especially challenging.
8 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Figure 3. Part of the lichen collection at the Olympic National Park curatorial facility.
Results 9
Tabl
e 1.
Mos
s, liv
erw
ort,
horn
wor
t and
lich
en sp
ecie
s rec
orde
d fr
om th
e O
lym
pic
Peni
nsul
a
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
HO
RN
WO
RTS
Ant
hoce
ros
fusi
form
is
72
147
-106
810
Ant
hoce
ros
punc
tatu
s 2
103-
879
2
2 H
OR
NW
OR
T sp
ecie
s fou
nd o
n O
P (2
004)
2 H
OR
NW
OR
T sp
ecie
s on
Rho
ades
list
(199
7)0
00
00
00
01
00
00
00
20
01
0
2 H
OR
NW
OR
TS in
ON
P H
erba
rium
(OLY
M) (
1997
)N
umbe
r of H
ORN
WO
RT sp
ecie
s fou
nd in
eac
h m
ajor
ve
geta
tion
type
Num
ber o
f H
ORN
WO
RT sp
ecie
s f o
und
per s
ubstr
ate
cate
gory
LIC
HEN
SA
brot
hallu
s w
elw
itsch
iiO
P sp
ecim
en a
t BG
Her
bariu
m
Aca
rosp
ora
fusc
ata
Aca
rosp
ora
smar
agdu
la
Aht
iana
pal
lidul
a 1
141
41
13
11
12
OLY
M c
oll.
F. R
hoad
es33
5-16
2716
Aht
iana
sph
aero
spor
ella
3
Gle
w 1
998,
WN
HP
Jeffe
rson
1150
-203
22
Ale
ctor
ia im
shau
gii
111
11
120
2-18
0714
Ale
ctor
ia la
ta
1?
McC
une:
inte
rgr.
w A
LSA
1461
1
Ale
ctor
ia n
igric
ans
21
13
2G
lew
199
8, W
NH
P Cl
alla
m, J
effe
rson
1627
-203
26
Ale
ctor
ia o
chro
leuc
a 4
verif
. K. G
lew
1873
-203
24
Ale
ctor
ia s
arm
ento
sa s
ubsp
. sa
rmen
tosa
4133
52
115
49
11
13
34
54
117
OP
spec
imen
at B
G H
erba
rium
5-20
3210
2
Ale
ctor
ia s
arm
ento
sa s
ubsp
. ve
xilli
fera
11
31
1G
lew
199
868
5-20
326
Ale
ctor
ia v
anco
uver
ensi
s 18
35
21
117
-116
325
Alla
ntop
arm
elia
alp
icol
aG
lew
199
8
Am
andi
nea
punc
tata
Am
ygda
laria
pan
aeol
a
Arc
tom
ia in
terf
ixa
Art
honi
a ili
cina
Tø
nsbe
rg 1
995,
199
9
Art
honi
a le
ucop
ella
eaTø
nsbe
rg 1
998
Art
honi
a pr
uina
ta
OP
spec
imen
at B
G H
erba
rium
Art
honi
a ra
diat
a
14
Tabl
e 1.
Sp
ecie
s lis
t of m
osse
s, li
verw
orts
, hor
nwor
ts, l
iche
ns a
nd a
ssoc
iate
d ve
geta
tion
type
s an
d su
bstra
te in
Oly
mpi
c N
atio
nal P
ark,
Was
hing
ton.
(F
ootn
otes
on
page
52.
)
10 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Art
honi
a st
ella
ris
Tøns
berg
199
5, W
NH
P Cl
alla
m,
Jeffe
rso n
Art
hoth
eliu
m n
orve
gicu
m1
OP
spec
imen
at B
G H
erba
rium
51
Art
hoth
eliu
m s
pect
abile
det.
John
Vill
ella
1031
1
Art
hror
haph
is a
erug
inos
aO
P sp
ecim
en a
t BG
Her
bariu
m
Art
hror
haph
is c
itrin
ella
Asp
icili
a al
iena
Asp
icili
a ca
esio
cine
rea
Gle
w 1
998
Asp
icili
a ca
ndid
aBr
odo
et a
l. 20
01, G
lew
199
8
Asp
icili
a ci
nere
a
Asp
icili
a gi
bbos
a
Asp
icili
a la
evat
a
Bac
idia
arc
eutin
a
Bac
idia
het
eroc
hroa
OP
spec
imen
at B
G H
erba
rium
Bac
idia
sal
mon
eaO
P sp
ecim
en a
t BG
Her
bariu
m
Bac
idia
viri
difa
rinos
aTø
nsbe
rg 1
997
Bac
idin
a ar
nold
iana
OP
spec
imen
at B
G H
erba
rium
Bac
idin
a in
unda
ta
Bac
idin
a ph
acod
es
Bae
omyc
es r
ufus
4
11
OP
spec
imen
at B
G H
erba
rium
16-9
077
Bel
lem
erea
alp
ina
Bel
lem
erea
cin
ereo
rufe
scen
s G
lew
199
8
Bia
tora
cup
rea
Bia
tora
eff
lore
scen
s O
P sp
ecim
en a
t BG
Her
bariu
m
Bia
tora
fla
vopu
ncta
ta
Prin
tzen
& T
ønsb
erg
1999
(Tø
nsbe
rg
1 665
9 in
ON
P )B
iato
ra h
ypop
haea
Prin
tzen
& T
ønsb
erg
1999
Bia
tora
mei
ocar
pa v
ar.
taco
men
sis
Prin
tzen
& T
ønsb
erg
1999
Bia
tora
pau
siac
aO
P sp
ecim
en a
t BG
Her
bariu
m
Bia
tora
por
phyr
ospo
da
Tøns
berg
199
3
Bia
tora
sub
dupl
exO
P sp
ecim
en a
t BG
Her
bariu
m
Bia
tora
toe
nsbe
rgii
Prin
tzen
& T
ønsb
erg
1999
Bia
tora
vac
cini
icol
a Pr
intz
en &
Tøn
sber
g 19
99 (T
ønsb
erg
1 666
0 in
ON
P )
15
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 11
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Bia
tora
ver
nalis
Bro
doa
oroa
rctic
a 12
15
WN
HP
Clal
lam
1630
-203
28
Bry
onor
a sp
. B
. cas
tane
a sp
ecim
en a
t OLY
M, n
ot
verif
.B
ryor
ia b
icol
or
6ve
rif. R
. Ros
entre
tter
5-81
43
Bry
oria
cap
illar
is
262
16
11
15-
1271
23
Bry
oria
cha
lybe
iform
is
92
22
948-
2032
7
Bry
oria
fre
mon
tii
123
13
31
41
11
12
157-
1746
26
Bry
oria
fria
bilis
3
1W
NH
P Cl
alla
m79
-758
3
Bry
oria
fur
cella
ta
11
657-
1233
2
Bry
oria
fus
cesc
ens
492
14
111
23
21
641
9-19
5837
Bry
oria
gla
bra
161
27
11
123-
1623
16
Bry
oria
impl
exa
71
177
5-18
076
Bry
oria
lane
stris
O
LYM
spec
. misd
eter
m
Bry
oria
'mys
tery
spe
cies
' 6
11
51
180-
1627
9
Bry
oria
niti
dula
22
spec
imen
s nee
d ve
rific
atio
n18
07-2
032
2
Bry
oria
pik
ei
WN
HP
Clal
lam
Bry
oria
pse
udoc
apill
aris
L.
Gei
ser 2
000
Bry
oria
pse
udof
usce
scen
s 25
25
248
-162
018
Bry
oria
sub
cana
L.
Gei
ser 2
000
Bry
oria
ten
uis
2ID
qui
te fi
rm b
ut w
ill b
e se
nt to
Bro
do16
-825
2
Bry
oria
tor
tuos
a 3
verif
. T. T
ønsb
erg,
WN
HP
Clal
lam
79-4
242
Bry
oria
tric
hode
s 5
15-
624
4
Bry
oria
tric
hode
s su
bsp.
am
eric
ana
17-2
82
Bry
oria
tric
hode
s su
bsp.
tric
hode
s
Bue
llia
disc
iform
is
228
91
Bue
llia
erub
esce
ns
Tøns
berg
199
8
Bue
llia
geop
hila
Gle
w 1
998
Bue
llia
gris
eovi
rens
Tøns
berg
199
8
Bue
llia
mic
robo
la
Bue
llia
mur
iform
isO
P sp
ecim
en a
t BG
Her
bariu
m
Bue
llia
oida
lea
OLY
M c
oll.
Coo
ke
16
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
12 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Bue
llia
papi
llata
Bue
llia
still
ingi
ana
Bue
llia
turg
esce
ns
Bun
odop
horo
n m
elan
ocar
pum
3
21
verif
. B. M
cCun
e, O
P sp
ecim
en a
t BG
H
erba
ri um
8-16
74
Bys
solo
ma
mar
gina
tum
OP
spec
imen
at B
G H
erba
rium
Cal
iciu
m a
biet
inum
O
LYM
col
l F. R
hoad
es
Cal
iciu
m g
lauc
ellu
m
OLY
M c
oll F
. Rho
ades
Cal
iciu
m le
ntic
ular
e1
182
1
Cal
iciu
m s
alic
inum
Cal
iciu
m v
iride
2
11
123-
624
3
Cal
opla
ca a
mm
iosp
ilaG
lew
199
8
Cal
opla
ca a
troa
lba
Cal
opla
ca a
tros
angu
inea
Tø
nsbe
rg 1
999c
Cal
opla
ca b
olac
ina
Cal
opla
ca c
erin
a
Cal
opla
ca c
f. b
orea
lisO
NP
spec
imen
det
. C. W
etm
ore
2004
Cal
opla
ca c
itrin
a O
P sp
ecim
en a
t BG
Her
bariu
m
Cal
opla
ca c
renu
laria
Cal
opla
ca d
ispe
rsa
Gle
w 1
998
Cal
opla
ca e
pith
allin
a G
lew
199
8
Cal
opla
ca f
erru
gine
a
Cal
opla
ca f
lavo
gran
ulos
a
Cal
opla
ca f
lavo
rube
scen
s
Cal
opla
ca h
oloc
arpa
gro
upO
NP
spec
imen
in B
G H
erba
rium
sp
ecim
enco
nf.C
.Wet
mor
e 2 0
04C
alop
laca
jung
erm
anni
ae
Cal
opla
ca la
mpr
oche
ilaG
lew
199
8
Cal
opla
ca li
toric
ola
OP
spec
imen
at B
G H
erba
rium
Cal
opla
ca lu
teom
inia
var
. lu
teom
inia
Cal
opla
ca m
arin
a su
bsp.
am
eric
ana
Cal
opla
ca r
osei
Cal
opla
ca s
axic
ola
Gle
w 1
998
17
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 13
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Cal
opla
ca s
inap
ispe
rma
Cal
opla
ca s
oroc
arpa
O
NP
spec
imen
in B
G H
erba
rium
sp
ecim
e nco
nf.C
.Wet
mo r
e 20
0 4C
alop
laca
tiro
liens
is
Gle
w 1
998
Can
dela
ria c
onco
lor
193
1
Can
dela
riella
aur
ella
G
lew
199
8
Can
dela
riella
ref
lexa
OP
spec
imen
at B
G H
erba
rium
Can
dela
riella
ter
rigen
aG
lew
199
8
Can
dela
riella
vite
llina
G
lew
199
8
Can
dela
riella
xan
thos
tigm
aO
P sp
ecim
en a
t BG
Her
bariu
m
Cat
apyr
eniu
m c
iner
eum
Cat
apyr
eniu
m d
aeda
lum
1
verif
. B. M
cCun
e16
271
Cat
illar
ia f
ranc
isca
na
Cat
inar
ia a
trop
urpu
rea
Cav
ernu
laria
hul
teni
i 53
51
9O
P sp
ecim
en a
t BG
Her
bariu
m5-
1307
42
Cav
ernu
laria
loph
yrea
1
152-
939
24
Cet
raria
acu
leat
a 3
310
516
30-2
032
7
Cet
raria
are
naria
Cet
raria
eric
etor
um
24
72
1167
-203
210
Cet
raria
eric
etor
um s
ubsp
. er
icet
orum
Cet
raria
eric
etor
um s
ubsp
. re
ticul
ata
Cet
raria
isla
ndic
a su
bsp.
cris
pifo
rmis
Cet
raria
isla
ndic
a su
bsp.
isla
ndic
a1
313
414
82-2
032
10
Cet
raria
laev
igat
a
Cet
raria
mur
icat
a 2
61
94
1238
-203
28
Cet
relia
cet
rario
ides
5
12
5-28
97
Cha
enot
heca
fer
rugi
nea
OP
spec
imen
at B
G H
erba
rium
Cha
enot
heca
fur
fura
cea
31
2O
P sp
ecim
en a
t BG
Her
bariu
m5-
1271
4
Chr
ysot
hrix
can
dela
ris
21
11
Tøns
berg
199
9c5-
707
8
Chr
ysot
hrix
chl
orin
a 1
21
130-
822
3
Chr
ysot
hrix
chr
ysop
htha
lma
OP
spec
imen
at B
G H
erba
rium
Chr
ysot
hrix
gra
nulo
saO
P sp
ecim
en a
t BG
Her
bariu
m
18
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
14 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Cla
dina
arb
uscu
la
Gle
w 1
998
Cla
dina
arb
uscu
la
var.
ber
inge
riana
Brod
o et
al.
2001
Cla
dina
miti
s 2
21
95
212-
1978
12
Cla
dina
por
tent
osa
11
146
-778
5
Cla
dina
por
tent
osa
subs
p. p
acifi
ca1
16-1
763
Cla
dina
ran
gife
rina
22
112
117
6-12
3817
Cla
dina
ste
llaris
Cla
doni
a al
boni
gra
21
128
-127
13
Cla
doni
a ar
tuat
a
Cla
doni
a as
ahin
ae
Cla
doni
a ba
cilli
form
is
11
182-
685
2
Cla
doni
a be
llidi
flora
5
511
1821
25
13
23
21
42
85-
1746
48
Cla
doni
a ca
riosa
1
11
Gle
w 1
998
1511
-195
83
Cla
doni
a ca
rneo
la
21
1911
13
31
21
31
628
-167
830
Cla
doni
a ce
note
a 2
115
9-77
83
Cla
doni
a ce
rvic
orni
s su
bsp.
cer
vico
rnis
11
13
948-
1886
5
Cla
doni
a ce
rvic
orni
s su
bsp.
ver
ticill
ata
113
481
Cla
doni
a ch
loro
phae
a 5
52
615
21
72
21
35-
1745
22
Cla
doni
a co
ccife
ra
111
60-1
930
2
Cla
doni
a co
nioc
raea
1
610
23
22
11
11
31
413
4-16
2019
Cla
doni
a co
rnut
a su
bsp.
cor
nuta
15-
182
2
Cla
doni
a cr
ispa
ta v
ar.
cris
pata
13
22
116
-389
5
Cla
doni
a cy
anip
es
22
637
1
Cla
doni
a de
cort
icat
a 11
671
Cla
doni
a de
form
is
Cla
doni
a di
gita
ta
24
12
159-
634
6
Cla
doni
a di
mor
pha
Cla
doni
a ec
moc
yna
14
184
723-
2032
15
Cla
doni
a ec
moc
yna
subs
p. e
cmoc
yna
Cla
doni
a ec
moc
yna
subs
p. in
term
edia
12
182
5-14
374
Cla
doni
a ec
moc
yna
subs
p. o
ccid
enta
lis1
14
11
282
5-18
078
19
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 15
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Cla
doni
a fim
bria
ta
32
13
52
41
15-
1627
18
Cla
doni
a fu
rcat
a 3
111
11
176-
1167
14
Cla
doni
a gr
acili
s 1
21
Gle
w 1
998
1437
-174
64
Cla
doni
a gr
acili
s su
bsp.
elo
ngat
a16
271
Cla
doni
a gr
acili
s su
bsp.
tur
bina
ta1
1160
1
Cla
doni
a gr
acili
s su
bsp.
vul
nera
ta1
1807
1
Cla
doni
a gr
ayi
12
110
93-1
627
3
Cla
doni
a hu
mili
s
Cla
doni
a m
acile
nta
12
12
93-1
164
3
Cla
doni
a m
acile
nta
var.
bac
illar
is
Cla
doni
a m
acro
phyl
la
Cla
doni
a m
acro
phyl
lode
s 4
22
1325
-195
86
Cla
doni
a m
eroc
hlor
opha
ea
22
12
11
210-
1292
5
Cla
doni
a m
etac
oral
lifer
a 1
112
181
Cla
doni
a m
ultif
orm
is
116
-116
02
Cla
doni
a no
rveg
ica
214
14
13
11
4O
P sp
ecim
en a
t BG
Her
bariu
m,
T øn s
ber g
&G
owar
d 1 9
92,W
NH
P19
6-12
9213
Cla
doni
a no
voch
loro
phae
a 1
219
50-1
978
2
Cla
doni
a oc
hroc
hlor
a 1
110
251
56
24
21
31
27
5-16
2928
Cla
doni
a ph
yllo
phor
a 1
1167
1
Cla
doni
a pl
euro
ta
11
162
4-11
602
Cla
doni
a po
cillu
m
13
1167
-195
84
Cla
doni
a py
xida
ta
35
11
41
31
15-
1978
18
Cla
doni
a re
i 1
11
637-
1292
2
Cla
doni
a sc
abriu
scul
a 4
19
119
12
16
12
24
25-
1958
27
Cla
doni
a sc
hofie
ldii
11
1745
-195
02
Cla
doni
a si
ngul
aris
1
822-
1348
2
Cla
doni
a sp
.1
23
11
219
-564
5
Cla
doni
a sq
uam
osa
var.
squ
amos
a2
24
1715
32
21
12
42
0-14
3730
Cla
doni
a sq
uam
osa
var.
sub
squa
mos
a1
718
11
11
53
71
13
15-
1164
28
Cla
doni
a st
ricta
Cla
doni
a su
bula
ta
11
1002
1
20
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
16 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Cla
doni
a su
lphu
rina
14
11
723-
1476
6
Cla
doni
a sy
mph
ycar
pa
116
291
Cla
doni
a tr
ansc
ende
ns
1124
21
12
22
74
34
22
5-12
6129
Cla
doni
a tu
rgid
a
Cla
doni
a um
bric
ola
1424
15
32
21
52
24
5-15
1131
Cla
doni
a un
cial
is
Cla
doni
a ve
rruc
ulos
a 2
12
13
1134
-151
14
Clio
stom
um f
lavi
dulu
mTø
nsbe
rg 1
997
Clio
stom
um g
riffit
hii
Tøns
berg
199
9c
Clio
stom
um le
pros
umO
P sp
ecim
en a
t BG
Her
bariu
m
Coc
cotr
ema
poci
llariu
m
2Tø
nsbe
rg 1
998
WN
HP
Clal
lam
51
Col
lem
a fe
cund
um
12
1
Col
lem
a fu
rfur
aceu
m
OP
spec
imen
at B
G H
erba
rium
Col
lem
a fu
scov
irens
1
cf.
(M. H
utte
n sp
ecim
en)
1624
1
Col
lem
a gl
ebul
entu
m
116
301
Col
lem
a oc
culta
tum
O
P sp
ecim
en a
t BG
Her
bariu
m
Col
lem
a sp
. 1
prel
imin
ary,
to b
e ve
rifie
d94
81
Col
lem
a un
dula
tum
var
. gr
anul
osum
1pr
elim
inar
y, to
be
verif
ied
2032
1
Cor
nicu
laria
nor
moe
rica
114
WN
HP
Clal
lam
1630
-203
29
Cyp
heliu
m in
quin
ans
11
123-
778
3
Cyp
heliu
m lu
cidu
m
Cyp
heliu
m p
inic
ola
Cyp
heliu
m t
igill
are
Cys
toco
leus
ebe
neus
O
P sp
ecim
en a
t BG
Her
bariu
m,
WN
HP
Gr a
ys H
arbo
rD
acty
losp
ora
para
sitic
aO
P sp
ecim
en a
t BG
Her
bariu
m
Den
dris
coca
ulon
sp.
2ve
rif. C
. Der
r5-
317
2
Der
mat
ocar
pon
inte
stin
iform
e 1
1958
1
Der
mat
ocar
pon
lurid
um
WN
HP
Clal
lam
, Jef
fers
on, S
&M
do
cum
e nts
Der
mat
ocar
pon
min
iatu
m
21
182
2-20
324
Der
mat
ocar
pon
mou
linsi
i 94
81
Der
mat
ocar
pon
retic
ulat
um2
1539
-203
25
21
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 17
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Der
mat
ocar
pon
rivul
orum
41
1348
-195
06
Dim
erel
la lu
tea
OP
spec
imen
at B
G H
erba
rium
Dim
erel
la p
inet
i 1
1O
P sp
ecim
en a
t BG
Her
bariu
m10
441
Dip
losc
hist
es s
crup
osus
Dip
loto
mm
a al
boat
rum
Dip
loto
mm
a pe
nich
rum
End
ocar
pon
pusi
llum
118
861
End
ococ
cus
apic
iicol
aO
P sp
ecim
en a
t BG
Her
bariu
m
Erio
derm
a so
redi
atum
3
verif
. B. M
cCun
e, O
P sp
ecim
en a
t BG
H
erba
ri um
5-28
93
Ess
linge
riana
idah
oens
is
71
126
5-15
258
Euo
psis
gra
natin
a
Eve
rnia
pru
nast
ri 8
12
30-
419
12
Fel
lhan
era
bout
eille
i6
OP
spec
imen
at B
G H
erba
rium
7-18
03
Fel
lhan
erop
sis
vezd
aeO
P sp
ecim
en a
t BG
Her
bariu
m
Fis
surin
a in
sidi
osa
Fla
voce
trar
ia c
ucul
lata
2
25
316
30-2
032
8
Fla
voce
trar
ia n
ival
is
21
83
1807
-203
27
Fla
vopu
ncte
lia s
ored
ica
Fus
copa
nnar
ia la
cera
tula
23
11
11
verif
. B. M
cCun
e5-
861
5
Fus
copa
nnar
ia le
ucop
haea
O
P sp
ecim
en s.
l. at
BG
Her
bariu
m
Fus
copa
nnar
ia le
ucos
ticto
ides
2
171
33
21
OP
spec
imen
at B
G H
erba
rium
5-86
515
Fus
copa
nnar
ia m
ariti
ma
Not
e ou
r mat
eria
l nee
ds to
be
be
r ede
term
ined
Fus
copa
nnar
ia m
edite
rran
ea
11
Not
e ou
r mat
eria
l nee
ds to
be
be
r ede
term
ined
389
1
Fus
copa
nnar
ia p
acifi
caO
P sp
ecim
en a
t BG
Her
bariu
m, o
urs:
r e- d
eter
min
ati o
n s r e
quir e
d5
1
Fus
copa
nnar
ia p
raet
erm
issa
1
11
11
574-
1807
5
Fus
copa
nnar
ia s
aubi
netii
2
193
11
16
re-d
eter
min
atio
ns re
quire
d18
3-11
8125
Gra
phis
ele
gans
O
P sp
ecim
en a
t BG
Her
bariu
m
Gra
phis
scr
ipta
1
1O
P sp
ecim
en a
t BG
Her
bariu
m12
3-28
94
Gya
lideo
psis
ana
stom
osan
s Tø
nsbe
rg 1
995
Gya
lideo
psis
mus
cico
laTø
nsbe
rg a
nd H
enss
en 1
999
Gya
lideo
psis
pic
eico
la
1Tø
nsbe
rg 1
995
465
1
22
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
18 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Hae
mat
omm
a pe
rsoo
nii
spec
imen
at O
NP
herb
ariu
m, n
ot
verif
ied
Hal
ecan
ia v
iride
scen
s Tø
nsbe
rg 1
998,
WN
HP
Clal
lam
Het
erod
erm
ia le
ucom
ela
L. G
eise
r 200
0
Hyd
roth
yria
ven
osa
1
Hyp
ocen
omyc
e ca
stan
eoci
nere
a 3
71
22
116
-162
410
Hyp
ocen
omyc
e fr
iesi
i 2
900-
1624
2
Hyp
ocen
omyc
e le
ucoc
occa
1
11
tent
ativ
e O
NP
spec
.15
111
Hyp
ocen
omyc
e sc
alar
is
22
12
618-
1629
5
Hyp
ogym
nia
apin
nata
64
11
211
31
33
33
16
21
55-
1627
67
Hyp
ogym
nia
aust
erod
es
11
Gle
w 1
998
1958
1
Hyp
ogym
nia
bitt
eri
Hyp
ogym
nia
dupl
icat
a 1
551
11
23
4ve
rif. C
. Der
r, W
NH
P Cl
all,
Jeff,
G
rays
H.
8-16
2748
Hyp
ogym
nia
ente
rom
orph
a 1
1110
51
144
83
43
142
48
411
OP
spec
imen
at B
G H
erba
rium
0-16
2981
Hyp
ogym
nia
imsh
augi
i 2
494
21
121
22
33
16
123-
2032
49
Hyp
ogym
nia
inac
tiva
126
21
25-
1261
33
Hyp
ogym
nia
met
aphy
sode
s 16
22
16
21
13
verif
. B. M
cCun
e61
8-16
2021
Hyp
ogym
nia
occi
dent
alis
17
21
24
41
317
7-18
8621
Hyp
ogym
nia
ocea
nica
5
11
11
190-
822
5
Hyp
ogym
nia
phys
odes
1
383
11
13
26
11
32
22
0-18
0751
Hyp
ogym
nia
rugo
sa
127
32
32
81
28
624-
2032
36
Hyp
ogym
nia
tubu
losa
9
10-
1299
14
Hyp
ogym
nia
vitt
ata
verif
y so
urce
s! W
NH
P G
rays
Har
bor
Hyp
otra
chyn
a si
nuos
a 9
31
OP
spec
imen
at B
G H
erba
rium
5-82
916
Icm
adop
hila
eric
etor
um
116
12
21
12
11
55-
1271
28
Iona
spis
lacu
stris
Jape
wia
sub
aurif
era
OP
spec
imen
at B
G H
erba
rium
Kae
rnef
eltia
cal
iforn
ica
11
verif
. B. M
cCun
e, W
NH
P O
P da
ta in
er
r or ?
51
Kae
rnef
eltia
mer
rillii
17
11
12
313
0-20
3218
Koe
rber
ia s
onom
ensi
s 2
McC
une
says
our
Ves
terg
reno
psis
i sidi
ata
are
KO
SO14
78-1
624
2
Leca
nact
is a
biet
ina
Leca
nia
cyrt
ella
23
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 19
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Leca
nia
naeg
elii
Leca
nora
alb
ella
var
. al
bella
Leca
nora
allo
phan
a
Leca
nora
bic
inct
aG
lew
199
8
Leca
nora
cad
ubria
e
Leca
nora
cen
isia
Leca
nora
cin
ereo
fusc
a va
r. c
iner
eofu
sca
OP
spec
imen
at B
G H
erba
rium
Leca
nora
circ
umbo
real
is
Leca
nora
dis
pers
a G
lew
199
8
Leca
nora
epi
bryo
n G
lew
199
8
Leca
nora
exp
alle
ns
OP
spec
imen
at B
G H
erba
rium
Leca
nora
far
inar
ia
1Tø
nsbe
rg 1
995,
199
928
91
Leca
nora
fus
cesc
ens
Leca
nora
hag
enii
Gle
w 1
998
Leca
nora
jam
esii
Tøns
berg
199
7, 1
999
Leca
nora
mal
aena
Gle
w 1
998
Leca
nora
mur
alis
Leca
nora
pac
ifica
Leca
nora
pol
ytro
pa
Gle
w 1
998
Leca
nora
pul
icar
is
15-
52
Leca
nora
rup
icol
a
Leca
nora
sym
mic
ta
Leca
nora
um
bros
a
Leca
nora
var
ia
Leca
nora
xyl
ophi
la
Leci
dea
atro
brun
nea
Gle
w 1
998
Leci
dea
casc
aden
sis
Gle
w 1
998
Leci
dea
furv
onig
rans
Leci
dea
fusc
oatr
a
Leci
dea
lapa
cida
G
lew
199
8
Leci
dea
prae
nubi
la
24
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
20 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Leci
dea
rose
otin
cta
OP
spec
imen
at B
G H
erba
rium
Leci
dea
tess
ella
ta
Gle
w 1
998
Leci
dea
wul
feni
iG
lew
199
8
Leci
della
ano
mal
oide
s
Leci
della
ela
eoch
rom
a Tø
nsbe
rg 1
998
Leci
della
eup
hore
a 1
91
1134
-116
73
Leci
della
stig
mat
ea
Leci
della
wul
feni
i
Leci
dom
a de
mis
sum
Gle
w 1
998
Leio
derm
a so
redi
atum
1
verif
. B. M
cCun
e, p
ers.
com
. T o
n sbe
r g 2
004
81
Lepr
aria
cac
umin
umG
lew
199
8
Lepr
aria
cae
sioa
lba
11
118
1-94
83
Lepr
aria
diff
usa
var.
diff
usa
Lepr
aria
ebu
rnea
OP
spec
imen
at B
G H
erba
rium
Lepr
aria
inca
na
21
11
134-
1218
4
Lepr
aria
lobi
fican
s14
51
23
12
11
OP
spec
imen
at B
G H
erba
rium
109-
1629
14
Lepr
aria
neg
lect
a G
lew
199
818
21
Lepr
aria
vou
auxi
i
Lepr
ocau
lon
mic
rosc
opic
um
Lepr
ocau
lon
suba
lbic
ans
36
Gle
w 1
998
1150
-203
27
Lept
ochi
dium
alb
ocili
atum
4
775-
1150
4
Lept
ogiu
m b
rebi
sson
ii 9
OP
spec
imen
at B
G H
erba
rium
5-31
76
Lept
ogiu
m c
alifo
rnic
um
21
11
120
8-10
024
Lept
ogiu
m c
orni
cula
tum
6
56
131
21
12
OP
spec
imen
at B
G H
erba
rium
0-11
6722
Lept
ogiu
m c
yane
scen
s 4
71
13
26
OP
spec
imen
at B
G H
erba
rium
5-12
3315
Lept
ogiu
m g
elat
inos
um
15-
165
2
Lept
ogiu
m li
chen
oide
s 8
11
12
130
7-16
3012
Lept
ogiu
m m
inut
issi
mum
294
8-20
322
Lept
ogiu
m p
laty
num
1
277
5-20
323
Lept
ogiu
m p
olyc
arpu
m
35
24
12
1O
P sp
ecim
en a
t BG
Her
bariu
m5-
1265
14
Lept
ogiu
m s
atur
ninu
m
118
071
25
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 21
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Lept
ogiu
m s
ubar
idum
77
81
Lept
ogiu
m t
enui
ssim
um
21
392-
1160
3
Lept
ogiu
m t
eret
iusc
ulum
3
12
182-
1000
7
Leth
aria
col
umbi
ana
21
OP
spec
imen
at B
G H
erba
rium
1590
-203
26
Leth
aria
vul
pina
4
21
12
11
OP
spec
imen
at B
G H
erba
rium
248-
2032
16
Lich
enod
iplis
leca
noric
ola
OP
spec
imen
at B
G H
erba
rium
Loba
ria h
allii
8
11
1O
P sp
ecim
en a
t BG
Her
bariu
m,
WN
HP
Jeffe
rso n
180-
778
7
Loba
ria li
nita
12
291
11
6ve
rif. C
. Der
r20
2-20
3240
Loba
ria o
rega
na
541
25
15
11
44
35
OP
spec
imen
at B
G H
erba
rium
5-10
9647
Loba
ria p
ulm
onar
ia
117
21
23
21
11
1O
P sp
ecim
en a
t BG
Her
bariu
m5-
1292
33
Loba
ria s
crob
icul
ata
112
11
OP
spec
imen
at B
G H
erba
rium
5-11
5015
Loba
ria s
ilvae
-vet
eris
Tøns
berg
199
8
Lopa
dium
dis
cifo
rme
1M
cCun
e 26
896
subm
itted
to O
LYM
51
Lopa
dium
pez
izoi
deum
Loxo
spor
a el
atin
a Tø
nsbe
rg 1
998
Loxo
spor
opsi
s co
ralli
fera
4
1O
P sp
ecim
en a
t BG
Her
bariu
m5-
814
6
Mas
salo
ngia
car
nosa
3
61
14
44
26-1
627
16
Meg
alar
ia b
rodo
ana
Ekm
an &
Tøn
sber
g 19
96
Meg
alar
ia p
ulve
rea
OP
spec
imen
at B
G H
erba
rium
Meg
aspo
ra v
erru
cosa
G
lew
199
8
Mel
anel
ia c
omm
ixta
1
1G
lew
199
811
811
Mel
anel
ia d
isju
ncta
1
Gle
w 1
998
1630
1
Mel
anel
ia e
lega
ntul
a 10
118
07-1
958
4
Mel
anel
ia e
xasp
erat
a
Mel
anel
ia e
xasp
erat
ula
1G
lew
199
818
071
Mel
anel
ia f
ulig
inos
a 1
61
11
25-
1624
6
Mel
anel
ia h
epat
izon
7
16
Gle
w 1
998
1347
-203
28
Mel
anel
ia in
fum
ata
3G
lew
199
818
07-2
032
2
Mel
anel
ia m
ultis
pora
G
eise
r 200
0 da
ta
Mel
anel
ia p
anni
form
is
11
Gle
w 1
998
1403
-195
02
Mel
anel
ia s
ored
iata
2
Gle
w 1
998
1059
-180
72
26
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
22 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Mel
anel
ia s
tygi
a 6
2G
lew
199
818
07-2
032
6
Mel
anel
ia s
ubar
gent
ifera
Mel
anel
ia s
ubau
rifer
a 2
15-
778
3
Mel
anel
ia s
ubel
egan
tula
1
21
948-
1807
3
Mel
anel
ia s
ubol
ivac
ea
31
11
134-
1620
3
Mel
anel
ia t
omin
ii 1
ON
P 33
5 ne
eds w
ork,
it m
ay b
e to
o co
nve x
for M
ETO
1807
1
Men
egaz
zia
subs
imili
sBj
erke
, J.W
. 200
3
Men
egaz
zia
tere
brat
a 12
11
45
OP
spec
imen
at B
G H
erba
rium
, our
s: re
dete
rmin
atio
ns n
eede
d5-
317
16
Mic
area
cin
erea
OP
spec
imen
at B
G H
erba
rium
Mic
area
mic
roco
cca
OP
spec
imen
at B
G H
erba
rium
Mic
area
myr
ioca
rpa
Tøns
berg
199
9a
Mic
area
pel
ioca
rpa
OP
spec
imen
at B
G H
erba
rium
Mic
area
pra
sina
in
nee
d of
revi
sion:
mos
t, if
not a
ll r e
pres
ent M
. mi c
r oco
cca
(per
s.co
m.
Mic
area
pra
sine
llaO
P sp
ecim
en a
t BG
Her
bariu
m
Mic
area
syn
othe
oide
sTø
nsbe
rg, T
. and
B. C
oppi
ns 2
000
Mic
area
xan
thon
ica
1Co
ppin
s & T
ønsb
erg
2001
51
Mic
roly
chnu
s ep
icor
ticis
1
OP
spec
imen
at B
G H
erba
rium
, W
NH
P Cl
all a
m15
71
Mul
ticla
vula
muc
ida
31
165-
877
4
Mul
ticla
vula
ver
nalis
3
103-
1134
3
Myc
obili
mbi
a be
reng
eria
na1
179
1
Myc
obili
mbi
a sa
bule
toru
mG
lew
199
8
Myc
obla
stus
aff
inis
O
P sp
ecim
en a
t BG
Her
bariu
m, G
lew
19
98,
Myc
obla
stus
alp
inus
Myc
obla
stus
cae
sius
2Tø
nsbe
rg 1
999c
157
1
Myc
obla
stus
fuc
atus
Tø
nsbe
rg 1
3956
, Tøn
sber
g 13
765
NO
T in
ON
P ,bu
ton
OP
Myc
obla
stus
san
guin
ariu
s 15
31
42
11
12
OP
spec
imen
at B
G H
erba
rium
5-15
1122
Myc
opor
um a
ntec
elle
nsTø
nsbe
rg 1
998
Neo
fusc
elia
ver
rucu
lifer
a 1
1624
1
Nep
hrom
a be
llum
1
262
41
11
11
verif
. C. D
err
26-1
163
22
Nep
hrom
a he
lvet
icum
1
232
11
14
42
2O
P sp
ecim
en a
t BG
Her
bariu
m13
4-10
9328
Nep
hrom
a la
evig
atum
6
12
21
3O
P sp
ecim
en a
t BG
Her
bariu
m93
-473
12
27
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 23
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Nep
hrom
a oc
cultu
m
91
12
21
verif
. C. D
err
441-
1164
10
Nep
hrom
a pa
rile
310
201
42
31
22
118
3-17
4540
Nep
hrom
a re
supi
natu
m
326
11
13
11
17
1O
P sp
ecim
en a
t BG
Her
bariu
m93
-162
731
Nie
bla
ceph
alot
a 1
verif
. B. M
cCun
e, O
P sp
ecim
en a
t BG
H
erba
ri um
21
Nod
obry
oria
abb
revi
ata
32
1G
lew
199
815
11-2
032
5
Nod
obry
oria
ore
gana
1
551
14
52
222
12
19
335-
2032
47
Nod
obry
oria
sub
dive
rgen
s 3
11
215
11-2
032
5
Nor
man
dina
pul
chel
la
69
13
OP
spec
imen
at B
G H
erba
rium
5-11
349
Och
role
chia
and
rogy
na
1Tø
nsbe
rg 1
998
1629
1
Och
role
chia
far
inac
ea
11
spec
imen
s, ve
rific
atio
n re
ques
ted
389-
1807
2
Och
role
chia
inae
quat
ula
Och
role
chia
juve
nalis
Och
role
chia
laev
igat
a 6
11
11
134-
1271
7
Och
role
chia
ore
gone
nsis
376
22
14
11
12
23
5-19
5034
Och
role
chia
sub
atha
llina
Och
role
chia
sub
palle
scen
s 3
12
1147
-151
13
Och
role
chia
sza
tala
ënsi
s
Och
role
chia
tar
tare
a
Och
role
chia
ups
alie
nsis
1
62
Gle
w 1
998
419-
2032
7
Om
phal
ina
umbe
llife
ra
34
13
11
11
5-12
9920
Ope
grap
ha a
tra
Ope
grap
ha f
umos
aTø
nsbe
rg 2
5900
, in
Tøns
berg
199
9c,
T øn s
ber g
199
7O
pegr
apha
pro
tube
rans
Ope
grap
ha s
ored
iifer
a Tø
nsbe
rg 1
998,
WN
HP
Clal
lam
Ope
grap
ha t
helo
trem
atis
Tøns
berg
199
7, 1
999
Ope
grap
ha v
aria
Oph
iopa
rma
lapp
onic
a O
LYM
det
. B. R
yan,
WN
HP
Clal
l, Je
f fO
rphn
iosp
ora
mor
iops
is
Pan
naria
rub
igin
osa
124
1
Par
mel
ia h
ygro
phila
1
267
55
38
102
64
15
32
40-
1930
63
Par
mel
ia o
mph
alod
es
219
30-1
958
2
28
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
24 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Par
mel
ia p
seud
osul
cata
7
21
11
113
4-11
6710
Par
mel
ia s
axat
ilis
153
21
21
11
17-2
032
20
Par
mel
ia s
quar
rosa
9
15-
1167
9
Par
mel
ia s
ulca
ta
447
83
87
22
61
31
61
5-16
2356
Par
mel
iella
cya
nole
pra
247
-162
44
Par
mel
iella
par
vula
1
231
2O
P sp
ecim
en a
t BG
Her
bariu
m5-
1292
18
Par
mel
iella
trip
toph
ylla
7
1O
P sp
ecim
en a
t BG
Her
bariu
m, O
NP
spec
. not
sepa
r ate
dpe
rf ect
l y,s
ome
182-
512
7
Par
mel
iops
is a
mbi
gua
92
21
13
21
93-2
032
16
Par
mel
iops
is h
yper
opta
47
61
43
49
22
15
42
37
93-1
950
58
Par
mot
rem
a ar
nold
ii 14
15-
919
Par
mot
rem
a ch
inen
se
24-
52
Par
mot
rem
a cr
initu
m
15
5-28
6
Par
mot
rem
a pr
aeso
redi
osum
Pel
tiger
a ap
htho
sa
21
182
5-11
473
Pel
tiger
a br
itann
ica
44
137
163
41
31
62
2O
P sp
ecim
en a
t BG
Her
bariu
m5-
1299
46
Pel
tiger
a ca
nina
4
1134
-180
74
Pel
tiger
a ci
nnam
omea
1
15-
574
2
Pel
tiger
a co
llina
5
233
13
21
16
1O
P sp
ecim
en a
t BG
Her
bariu
m5-
1630
33
Pel
tiger
a de
geni
i 1
1746
1
Pel
tiger
a di
dact
yla
31
13
511-
1978
7
Pel
tiger
a ho
rizon
talis
Pel
tiger
a kr
istin
sson
ii 1
1G
lew
199
812
331
Pel
tiger
a le
pido
phor
a G
lew
199
8, W
NH
P Cl
alla
m
Pel
tiger
a le
ucop
hleb
ia
41
110
21
22
11
OP
spec
imen
at B
G H
erba
rium
265-
1629
20
Pel
tiger
a m
alac
ea
11
31
312
33-1
978
5
Pel
tiger
a m
embr
anac
ea
31
511
143
24
61
21
22
12
5-12
9940
Pel
tiger
a ne
cker
i 2
21
11
15-
1509
8
Pel
tiger
a ne
opol
ydac
tyla
2
17
154
33
34
16
12
5-12
6136
Pel
tiger
a pa
cific
a 4
24
11
11
11
8-15
0919
Pel
tiger
a po
lyda
ctyl
on
14
11
11
25-
1261
11
Pel
tiger
a po
noje
nsis
3
24
32
778-
1978
11
29
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 25
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Pel
tiger
a pr
aete
xtat
a 1
81
61
31
11
240
9-18
0719
Pel
tiger
a re
tifov
eata
2
1886
1
Pel
tiger
a ru
fesc
ens
34
75
111
60-2
032
13
Pel
tiger
a sc
abro
sa
Pel
tiger
a ve
nosa
4
11
11
11
15-
1620
16
Per
tusa
ria a
mar
a 1
OP
spec
imen
at B
G H
erba
rium
5-79
3
Per
tusa
ria b
orea
lis
21
OP
spec
imen
at B
G H
erba
rium
182-
210
2
Per
tusa
ria c
ocod
esTø
nsbe
rg 1
999a
Per
tusa
ria g
lauc
omel
a O
P sp
ecim
en a
t BG
Her
bariu
m
Per
tusa
ria le
iopl
aca
116
1
Per
tusa
ria o
phth
alm
iza
1O
P sp
ecim
en a
t BG
Her
bariu
m16
-182
2
Per
tusa
ria p
upill
aris
1O
P sp
ecim
en a
t BG
Her
bariu
m15
71
Per
tusa
ria s
ubam
bige
ns
OP
spec
imen
at B
G H
erba
rium
Pha
cops
is o
xysp
ora
Pha
eoca
liciu
m c
urtis
ii
Pha
eogr
aphi
s sm
ithii
Tøns
berg
199
9a
Pha
eoph
ysci
a co
nstip
ata
Pha
eoph
ysci
a en
doco
ccin
ea
Pha
eoph
ysci
a or
bicu
laris
3
1807
-188
62
Pha
eoph
ysci
a sc
iast
ra
1G
lew
199
818
071
Pha
eorr
hiza
nim
bosa
Gle
w 1
998
Phl
yctis
arg
ena
OP
spec
imen
at B
G H
erba
rium
Phl
yctis
spe
irea
OP
spec
imen
at B
G H
erba
rium
Phy
scia
ads
cend
ens
17
31
2-19
08
Phy
scia
aip
olia
7
14
174
-778
9
Phy
scia
biz
iana
3
1630
-193
03
Phy
scia
cae
sia
4G
lew
199
826
-203
24
Phy
scia
cal
losa
1
cf. v
ery
poor
spec
imen
2032
1
Phy
scia
dim
idia
ta
1G
lew
199
894
81
Phy
scia
dub
ia
118
071
Phy
scia
lept
alea
30
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
26 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Phy
scia
pha
ea
1O
LYM
seas
tack
; P.
cas
cade
nsis?
261
Phy
scia
ten
ella
1
61
OP
spec
imen
at B
G H
erba
rium
0-13
45
Phy
scon
ia a
mer
ican
a 2
247
3-18
072
Phy
scon
ia d
eter
sa
Phy
scon
ia f
alla
xde
t. Sa
rah
Jova
n, 2
003
431
Phy
scon
ia m
usci
gena
8
443
-203
26
Pilo
phor
us a
cicu
laris
10
31
22
2O
P sp
ecim
en a
t BG
Her
bariu
m15
3-12
9225
Pilo
phor
us c
lava
tus
94
15
153-
1348
18
Pilo
phor
us n
igric
aulis
6
1ve
rif. C
. Der
r63
4-16
274
Pla
cops
is g
elid
a 70
21
Pla
cops
is la
mbi
i1
mos
t of P
laco
pis g
elid
a is
P.la
mbi
i ac
c. M
cCun
e37
4-87
74
Pla
cynt
hiel
la u
ligin
osa
Gle
w 1
998
Pla
tism
atia
gla
uca
192
61
74
514
22
37
21
73
214
0-19
3097
Pla
tism
atia
her
rei
601
53
68
43
14
31
83
27
5-15
1181
Pla
tism
atia
lacu
nosa
28
22
12
verif
. C. D
err,
OP
spec
imen
at B
G
He r
bariu
m3-
1012
32
Pla
tism
atia
nor
vegi
ca
363
64
11
21
11
53
65-
1307
54
Pla
tism
atia
ste
noph
ylla
282
9-10
123
Ple
opsi
dium
chl
orop
hanu
m
118
071
Ple
opsi
dium
fla
vum
1ve
rif.
K. G
lew
1958
1
Pol
ychi
dium
Aff.
con
tort
um39
21
22
OP
spec
imen
at B
G H
erba
rium
5-10
1231
Pol
ychi
dium
mus
cico
la
16
11
138
9-16
278
Por
ina
lept
alea
Tøns
berg
199
9c
Por
ina
radi
cico
laM
cCar
thy
and
Tøns
berg
199
8
Por
pidi
a ca
rlotti
ana
Por
pidi
a co
ntra
poen
da
Por
pidi
a cr
ustu
lata
Por
pidi
a th
omso
nii
Pro
topa
nnar
ia p
eziz
oide
s8
64
35
31
37
Gle
w 1
998
5-16
9821
Pro
topa
rmel
ia b
adia
Pse
udep
hebe
min
uscu
la
145
948-
2032
9
Pse
udep
hebe
pub
esce
ns
3111
948-
2032
12
31
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 27
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Pse
udoc
yphe
llaria
ano
mal
a23
11
51
11
31
OP
spec
imen
at B
G H
erba
rium
5-90
730
Pse
udoc
yphe
llaria
ant
hras
pis
191
31
3O
P sp
ecim
en a
t BG
Her
bariu
m5-
822
17
Pse
udoc
yphe
llaria
cro
cata
29
11
45
verif
. C. D
err,
OP
spec
imen
at B
G
He r
bariu
m74
-104
825
Pse
udoc
yphe
llaria
mal
lota
20
11
12
2ve
rif. B
. McC
une,
Tøn
sber
g 19
99b
177-
907
17
Pse
udoc
yphe
llaria
per
petu
aU
SFS,
ver
if. C
. Der
r, pe
rs. c
om. D
. M
cCon
nal
Pse
udoc
yphe
llaria
rai
nier
ensi
s 21
11
11
12
verif
. C. D
err
202-
867
24
Psi
lole
chia
luci
daO
P sp
ecim
en a
t BG
Her
bariu
m
Pso
ra d
ecip
iens
OP
spec
imen
at B
G H
erba
rium
Pso
ra n
ippo
nica
3
571-
2032
3
Pso
rom
a hy
pnor
um
412
71
178
32
56
182-
2032
26
Pso
rula
ruf
onig
ra
Pyr
enul
a ac
utis
pora
OP
spec
imen
at B
G H
erba
rium
Pyr
enul
a la
evig
ata
Pyr
enul
a oc
cide
ntal
is
1M
cCun
e 26
900
subm
itted
to O
LYM
, O
Psp
ecim
enat
BG
Her
bari u
m5
1
Pyr
rhos
pora
cin
naba
rina
61
21
Mos
t (al
l?) O
P PY
CI re
cord
s con
cern
PY
GO
182-
1511
8
Pyr
rhos
pora
ela
bens
Pyr
rhos
pora
gow
ardi
ana
62
1T.
Spr
ibill
e ve
rifie
d se
vera
l OP
coll e
cti o
n s18
1-11
674
Pyr
rhos
pora
que
rnea
L.
Gei
ser 2
000,
WN
HP
Clal
lam
, OP
spec
imen
at B
G H
erba
ri um
Pyr
rhos
pora
sub
cinn
abar
ina
pers
. com
. Tøn
sber
g 20
03, W
NH
P C
lalla
mP
yrrh
ospo
ra v
aria
ns
Ram
alin
a di
lace
rata
6
1O
P sp
ecim
en a
t BG
Her
bariu
m91
-778
7
Ram
alin
a fa
rinac
ea
132
45
25-
335
19
Ram
alin
a m
enzi
esii
4W
NH
P Cl
alla
m, G
rays
Har
bor,
OP
spec
ime n
at B
G H
e rba
rium
4-69
5
Ram
alin
a po
llina
ria
L. G
eise
r 200
0, W
NH
P Cl
alla
m
Ram
alin
a ro
esle
ri 9
5-79
9
Ram
alin
a sp
. 2
21
0-2
2
Ram
alin
a su
blep
toca
rpha
4
6-43
2
Ram
alin
a th
raus
ta
22
113
0-19
02
Rhi
zoca
rpon
bad
ioat
rum
Rhi
zoca
rpon
bol
ande
ri G
lew
199
8
Rhi
zoca
rpon
cin
ereo
vire
ns
32
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
28 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Rhi
zoca
rpon
con
cent
ricum
Rhi
zoca
rpon
dis
poru
m
Gle
w 1
998
Rhi
zoca
rpon
gem
inat
um
Rhi
zoca
rpon
geo
grap
hicu
m
41
113
47-1
886
7
Rhi
zoca
rpon
gra
nde
Rhi
zoca
rpon
hoc
hste
tter
i
Rhi
zoca
rpon
leca
norin
um
Gle
w 1
998
Rhi
zoca
rpon
obs
cura
tum
Rhi
zoca
rpon
oed
eri
Rhi
zoca
rpon
rip
ariu
m
Rhi
zopl
aca
chry
sole
uca
2G
lew
199
813
16-1
627
2
Rhi
zopl
aca
mel
anop
htha
lma
132
Gle
w 1
998
1624
-203
28
Rim
ular
ia in
sula
risG
lew
199
8
Rin
odin
a di
sjun
cta
Tøns
berg
199
8
Rin
odin
a ha
llii
OP
spec
imen
at B
G H
erba
rium
Rin
odin
a m
niar
aea
Rin
odin
a st
ictic
a Sh
eard
& T
ønsb
erg
1995
Rop
alos
pora
viri
dis
Tøns
berg
199
8
Sag
iole
chia
rhe
xobl
epha
ra
San
tess
onie
lla g
risea
Tøns
berg
& H
enss
en 1
999
Sch
aere
ria c
ortic
ola
Tøns
berg
199
5
Sco
licio
spor
um s
arot
ham
niO
P sp
ecim
en a
t BG
Her
bariu
m
Sky
ttea
caes
iiO
P sp
ecim
en a
t BG
Her
bariu
m
Sky
ttea
leca
nora
eO
P sp
ecim
en a
t BG
Her
bariu
m
Sol
orin
a cr
ocea
9
11
1325
-193
08
Sol
orin
a sa
ccat
a
Sph
aero
phor
us g
lobo
sus
260
72
63
44
23
35
23
62
16
5-17
4786
Sph
aero
phor
us g
lobo
sus
var.
gra
cilis
169
-153
3
Spi
lone
ma
sp.1
1
1de
t. M
cCun
e, se
vera
l spe
cim
ens
177-
270
3
Spi
lone
mel
la a
mer
ican
a1
Hen
ssen
& T
ønsb
erg
2000
, typ
e lo
c.
in H
oh53
1
Spo
rast
atia
tes
tudi
nea
Gle
w 1
998
33
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 29
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Sta
urot
hele
are
olat
a
Sta
urot
hele
dru
mm
ondi
iG
lew
199
8
Sta
urot
hele
fis
sa
Ste
reoc
aulo
n al
pinu
m
21
1873
-203
23
Ste
reoc
aulo
n bo
tryo
sum
1
1630
1
Ste
reoc
aulo
n co
nden
satu
m
11
we
need
ref.
Spec
imen
s19
30-1
950
2
Ste
reoc
aulo
n gl
areo
sum
1
12
1G
lew
199
813
48-1
747
4
Ste
reoc
aulo
n gr
ande
2
219
50-1
978
3
Ste
reoc
aulo
n in
term
ediu
m
141
91
Ste
reoc
aulo
n pa
scha
le
21
392-
1930
4
Ste
reoc
aulo
n riv
ulor
um
Gle
w 1
998
Ste
reoc
aulo
n sa
saki
i var
. si
mpl
ex
Ste
reoc
aulo
n sa
saki
i var
. to
men
toso
ides
6
111
139
2-14
6117
Ste
reoc
aulo
n to
men
tosu
m
137
4-81
02
Ste
reoc
aulo
n ve
suvi
anum
1
we
need
ref.
Spec
imen
s11
631
Stic
ta b
eauv
oisi
i s.l.
202
33
verif
. C. D
err,
WN
HP
Jeffe
rson
177-
814
20
Stic
ta f
ulig
inos
a 5
341
13
21
12
62
OP
spec
imen
at B
G H
erba
rium
5-96
638
Stic
ta li
mba
ta
110
1O
P sp
ecim
en a
t BG
Her
bariu
m5-
711
11
Sul
caria
bad
ia
extin
ct?
Pet
erso
n et
al.
1998
, WN
HP
Cla
llam
Szc
zaw
insk
ia t
suga
e 1
det.
B. M
cCun
e, W
NH
P Cl
alla
m, O
P sp
ecim
enat
BG
Her
bari u
m53
1
Tep
hrom
ela
atra
O
P sp
ecim
en a
t BG
Her
bariu
m
Tha
mno
lia s
ubul
iform
is
33
18
312
38-2
032
11
Tha
mno
lia v
erm
icul
aris
3
66
353
9-20
3211
The
lom
ma
occi
dent
ale
det.
J. V
illel
la
The
lotr
ema
lepa
dinu
m
162
24
3O
P sp
ecim
en a
t BG
Her
bariu
m5-
1048
17
Tho
lurn
a di
ssim
ilis
E.Ti
sch
has O
P sp
ec.
Top
elio
psis
toe
nsbe
rgii
OLY
M c
oll.
Tøns
berg
Tra
pelia
cor
ticol
aO
P sp
ecim
en a
t BG
Her
bariu
m
Tra
pelio
psis
gel
atin
osa
Tra
pelio
psis
gra
nulo
sa
Gle
w 1
998
Tra
pelio
psis
pse
udog
ranu
losa
OP
spec
imen
at B
G H
erba
rium
34
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
30 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Tre
mel
la c
lado
niae
OP
spec
imen
at B
G H
erba
rium
Tre
mol
ecia
atr
ata
Gle
w 1
998
Tuc
kerm
anno
psis
chl
orop
hylla
1
514
51
310
11
25
33
42
OP
spec
imen
at B
G H
erba
rium
4-16
7859
Tuc
kerm
anno
psis
orb
ata
152
12
15
22
11
3-14
7630
Tuc
kerm
anno
psis
pla
typh
ylla
28
24
35
33
31
341
9-20
3228
Tuc
kerm
anno
psis
sub
alpi
na
316
63
52
1082
2-18
0722
Um
bilic
aria
ang
ulat
a 1
1W
NH
P Je
ffers
on62
41
Um
bilic
aria
arc
tica
41
OLY
M c
oll.
Ver
if. B
. Rya
n18
86-1
958
3
Um
bilic
aria
cyl
indr
ica
51
1150
-203
25
Um
bilic
aria
dec
ussa
ta
11
1ve
rif. B
. McC
une,
WN
HP
Cla
llam
1950
-195
82
Um
bilic
aria
deu
sta
413
47-1
807
3
Um
bilic
aria
hav
aasi
i 2
2ve
rif. B
. McC
une,
WN
HP
Cla
llam
948-
2032
5
Um
bilic
aria
hirs
uta
111
341
Um
bilic
aria
hyp
erbo
rea
281
994
8-20
3212
Um
bilic
aria
kra
sche
ninn
ikov
ii 6
1G
lew
199
8, W
NH
P Cl
alla
m, J
effe
rson
1886
-203
24
Um
bilic
aria
lam
bii
verif
. B. R
yan
Um
bilic
aria
lyng
ei
3ve
rif. B
. McC
une
1950
1
Um
bilic
aria
nyl
ande
riana
219
581
Um
bilic
aria
pha
ea
Um
bilic
aria
pol
yrrh
iza
194
81
Um
bilic
aria
pro
bosc
idea
3
4ve
rif. B
. McC
une,
WN
HP
Cla
llam
1930
-203
24
Um
bilic
aria
rig
ida
14ve
rif. B
. McC
une,
WN
HP
Cla
llam
, Je
ff er s
on18
07-2
032
5
Um
bilic
aria
sch
olan
deri
WN
HP
Mas
on
Um
bilic
aria
tor
refa
cta
146
Gle
w 1
998
728-
2032
9
Um
bilic
aria
vel
lea
119
821
Um
bilic
aria
virg
inis
4
2G
lew
199
813
25-2
032
5
Usn
ea c
aver
nosa
1
512
1
Usn
ea c
erat
ina
Usn
ea c
haet
opho
ra9
12
11
269
-127
19
Usn
ea c
ornu
ta
211
OP
spec
imen
at B
G H
erba
rium
5-69
8
Usn
ea d
iplo
typu
s 5
15-
452
9
35
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 31
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Usn
ea e
sper
antia
na2
69-7
42
Usn
ea f
ilipe
ndul
a 23
11
12
31
25-
1233
25
Usn
ea f
ragi
lesc
ens
Usn
ea f
ragi
lesc
ens
var.
mol
lis16
15-
1233
9
Usn
ea f
ulvo
reag
ens
281
Usn
ea g
labr
ata
31
4-26
54
Usn
ea g
labr
esce
ns s
ubsp
. gl
abre
la
Usn
ea h
espe
rina
7L.
Gei
ser 2
000
5-69
4
Usn
ea h
irta
Usn
ea la
ppon
ica
31
12-
134
6
Usn
ea lo
ngis
sim
a26
1W
NH
P Cl
alla
m, J
effe
rson
, Gra
ys
Har
bor ,
OP
spec
imen
at B
G5-
877
23
Usn
ea o
ccid
enta
lis3
5-28
93
Usn
ea p
acifi
cana
21
11
29
51
5-12
3312
Usn
ea r
ubic
unda
L.
Gei
ser 2
000
Usn
ea s
cabr
ata
144
12
21
69-1
271
11
Usn
ea s
cabr
ata
subs
p. n
ylan
deria
na77
81
Usn
ea s
ilesi
aca
82
8-62
46
Usn
ea s
phac
elat
a 2
2032
1
Usn
ea s
ubflo
ridan
a 13
11
22
15-
845
11
Usn
ea s
ubst
erili
s 5-
172
Usn
ea t
richo
dea
Usn
ea w
asm
uthi
i4
11
5-62
44
Usn
ea w
irthi
i 12
13
5-18
210
Ver
ruca
ria a
crot
ella
Ver
ruca
ria h
ydre
la
Ver
ruca
ria m
aura
Vul
pici
da c
anad
ensi
s 3
22
11
419-
1160
4
Vul
pici
da t
ilesi
i K
. Gle
w 1
998,
WN
HP
Clal
lam
Xan
thop
arm
elia
chl
oroc
hroa
Xan
thop
arm
elia
col
orad
oens
is
92
330-
2032
6
Xan
thop
arm
elia
cum
berla
ndia
1
61
36
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
32 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Xan
thop
arm
elia
wyo
min
gica
G
lew
199
8
Xan
thor
ia c
ande
laria
2
10-
263
Xan
thor
ia e
lega
ns
131
1G
lew
199
816
24-2
032
8
Xan
thor
ia f
alla
x 1
43-1
958
2
Xan
thor
ia p
olyc
arpa
4
12
5-13
47
Xan
thor
ia s
ored
iata
1
1950
1
Xyl
ogra
pha
hian
s
Xyl
ogra
pha
para
llela
Xyl
ogra
pha
vitil
igo
OP
spec
imen
at B
G H
erba
rium
711
LIC
HEN
spec
ies f
ound
on
OP
(200
4)
463
LIC
HEN
spec
ies o
n R
hoad
es li
st (1
997)
6879
7737
7556
7364
5849
8440
4357
3268
174
217
117
115
150
LIC
HEN
S in
ON
P H
erba
rium
(OLY
M) (
1997
)N
umbe
r of L
ICH
EN sp
ecie
s fou
nd in
eac
h m
ajor
ve
geta
tion
type
Num
ber o
f LIC
HEN
sp
ecie
s fou
nd p
er
subs
trate
cat
egor
y
LIV
ERW
OR
TSA
nast
roph
yllu
m m
inut
um1
21
21
1Cl
ark
& F
rye
1928
822-
1509
7
Ane
ura
ping
uis
27
47-1
747
11
Ant
helia
jura
tzka
na
216
85-1
747
3
Apo
met
zger
ia p
ubes
cens
2
141
793
-861
13
Ast
erel
la g
raci
lis
13
11
47-1
747
8
Ast
erel
la li
nden
berg
iana
4
21
12
571-
1747
7
Ast
erel
la s
acca
ta
113
481
Ath
alam
ia h
yalin
a 2
111
59-1
623
3
Bar
bilo
phoz
ia f
loer
kei
914
411
12
113
678-
2032
26
Bar
bilo
phoz
ia h
atch
eri
319
111
410
74
775-
2032
28
Bar
bilo
phoz
ia k
unze
ana
11
177
8-11
633
Bar
bilo
phoz
ia ly
copo
dioi
des
11
11
1H
ong
et a
l. 19
8925
4-16
786
Baz
zani
a am
bigu
a 1
11
134-
845
5
Baz
zani
a de
nuda
ta
113
225
210
26
5-82
226
Baz
zani
a tr
icre
nata
2
11
45-
167
6
Bla
sia
pusi
lla
121
11
25-
879
15
37
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 33
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Ble
phar
osto
ma
arac
hnoi
deum
3
11
Nor
ris 1
997
(spe
cim
en d
epos
ited
at
ON
P B .
trich
oph y
ll um
)20
2-10
934
Ble
phar
osto
ma
tric
hoph
yllu
m s
ubsp
. tr
icho
ph14
1314
8627
161
136
22
12
61
125
5-17
4574
Cal
ypog
eia
azur
ea
82
112
101
11
13
11
35-
1745
31
Cal
ypog
eia
fissa
9
22
146
11
42
13
36
5-17
4632
Cal
ypog
eia
inte
gris
tipul
a 1
102
11
25-
1096
10
Cal
ypog
eia
mue
lleria
na
212
132
83
13
19
48
115-
1437
48
Cal
ypog
eia
nees
iana
1
131
22
12
109-
1292
9
Cal
ypog
eia
spha
gnic
ola
23
5-28
74
Cal
ypog
eia
suec
ica
102
13
11
5-12
9210
Cep
halo
zia
ambi
gua
29
45-
1746
10
Cep
halo
zia
bicu
spid
ata
118
3101
155
210
119
73
61
28
5-16
2759
Cep
halo
zia
cate
nula
ta
Nor
ris 1
997
(Sh
ould
be
verif
ied,
few
re
cord
sin
WA
onH
ong'
s200
2W
AC
epha
lozi
a co
nniv
ens
25
192
12
21
28-1
164
17
Cep
halo
zia
leuc
anth
a
Cep
halo
zia
lunu
lifol
ia
210
106
49
411
33
19
101
84
205-
1307
64
Cep
halo
zia
mac
ouni
i
Cep
halo
zia
plen
icep
s 4
28-1
746
5
Cep
halo
ziel
la d
ivar
icat
a 5
73
135
24
21
12
31
15-
1745
34
Cep
halo
ziel
la d
ivar
icat
a va
r. s
cabr
a
Cep
halo
ziel
la r
ubel
la
Cep
halo
ziel
la s
tellu
lifer
a 15
71
Cep
halo
ziel
la t
urne
ri 4
13
289-
948
4
Chi
losc
yphu
s pa
llesc
ens
13
35
42
21
13
13
5-16
2323
Chi
losc
yphu
s po
lyan
thos
7
181
34
22
11
11
25-
1746
32
Chi
losc
yphu
s po
lyan
thos
var
. riv
ular
is
Con
ocep
halu
m c
onic
um
216
13
151
11
41
22
8-16
3032
Dip
loph
yllu
m a
lbic
ans
1832
35
94
34
4ve
rif. J
. Har
pel
5-10
9347
Dip
loph
yllu
m o
btus
ifoliu
m
155
13
26-1
745
18
Dip
loph
yllu
m p
licat
um
220
72
51
1ve
rif. J
. Har
pel
5-12
6118
Dip
loph
yllu
m ta
xifo
lium
13
363
27
62
13
427
Nor
ris 1
997
8-17
4741
Dou
inia
ova
ta
374
53
11
13
63
5-10
3161
38
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
34 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Fos
som
bron
ia f
oveo
lata
Fos
som
bron
ia lo
ngis
eta
Nor
ris 1
997
(Sh
ould
be
verif
ied,
s in
g le
sou r
ceo n
Ho n
g 's 2
002
WA
list
)F
rulla
nia
bola
nder
i 4
11
43-6
245
Fru
llani
a ca
lifor
nica
1
419
1
Fru
llani
a fr
anci
scan
a 23
13
2-12
311
Fru
llani
a ni
squa
llens
is
6160
83
55
36
814
17
193
13
5-10
1210
2
Geo
caly
x gr
aveo
lens
1
121
12
21
5-11
5017
Gym
noco
lea
infla
ta
21
10-1
674
Gym
nom
itrio
n co
ncin
natu
m
41
1Sc
hofie
ld 1
967
1348
-162
74
Gym
nom
itrio
n ob
tusu
m
18
32
1061
-150
97
Gyr
othy
ra u
nder
woo
dian
a 22
24
31
22
8-10
9624
Har
pant
hus
floto
vian
us
11
13
13
1093
-174
06
Her
bert
us a
dunc
us
17
33
verif
. J. H
arpe
l5-
176
12
Hyg
robi
ella
laxi
folia
Jam
eson
iella
aut
umna
lis
223
26
11
OLY
M c
oll.
Nor
ris5-
1079
6
Jung
erm
anni
a at
rovi
rens
5
43
5-10
4811
Jung
erm
anni
a co
nfer
tissi
ma
11
1ne
eds w
ork,
NEW
to W
A11
47-1
747
2
Jung
erm
anni
a ex
sert
ifolia
sub
sp.
cord
ifolia
142
571-
1747
11
Jung
erm
anni
a gr
acill
ima
15
1
Jung
erm
anni
a hy
alin
a 1
321
9-16
853
Jung
erm
anni
a le
iant
ha
21
14
12
15-
1461
11
Jung
erm
anni
a pu
mila
1
1
Jung
erm
anni
a ru
bra
242
21
5-70
216
Jung
erm
anni
a sp
haer
ocar
pa
113
911
Kur
zia
pauc
iflor
a 1
2de
t. Sc
hofie
ld a
nd H
arpe
l. N
OT
on
Hon
g 2 0
02 li
stw
i th h
epat
i cs.
8-13
43
Kur
zia
sylv
atic
a
Lepi
dozi
a fil
amen
tosa
Su
spec
t ver
ify!
Lepi
dozi
a re
ptan
s 8
602
64
12
26
73
68
25
5-13
0758
Loph
ocol
ea b
iden
tata
1
11
21
5-61
87
Loph
ocol
ea h
eter
ophy
lla
311
1322
78
46
1013
21
45
5-16
2027
Loph
ozia
asc
ende
ns
11
1620
1
39
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 35
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Loph
ozia
col
laris
1
113
91-1
590
2
Loph
ozia
exc
isa
120
321
Loph
ozia
gill
man
ii 21
1-13
912
Loph
ozia
het
eroc
olpo
s 1
21
21
11
128
-127
16
Loph
ozia
inci
sa
23
615
67
66
31
34
36
85-
1740
68
Loph
ozia
long
iden
s 3
11
177
8-16
274
Loph
ozia
long
iflor
a 1
13
644
88
318
12
614
Nor
ris 1
997
as L
. gut
tula
ta5-
1678
39
Loph
ozia
obt
usa
112
711
Loph
ozia
opa
cifo
lia
55
702-
1590
14
Loph
ozia
sud
etic
a 3
21
61
63
1096
-193
014
Loph
ozia
ven
tric
osa
47
85
31
81
15
778-
1740
20
Loph
ozia
wen
zelii
5
41
1810
845-
1747
14
Man
nia
sibi
rica
Nor
ris 1
997
Sho
uld
be v
erifi
ed a
s th
i sw
oul d
be n
ewto
WA
(not
onM
arch
antia
pol
ymor
pha
12
11
247
-174
79
Mar
supe
lla b
revi
ssim
a 1
31
31
1261
-150
94
Mar
supe
lla c
omm
utat
a
Mar
supe
lla c
onde
nsat
a 2
1348
-162
72
Mar
supe
lla e
mar
gina
ta
421
13
23
76-1
348
20
Mar
supe
lla e
mar
gina
ta v
ar.
aqua
tica
1cf
. onl
y fra
gmen
ts, r
evisi
t!13
48-1
685
2
Mar
supe
lla s
pars
ifolia
H
ong
et a
l. 19
89
Mar
supe
lla s
phac
elat
a 3
51
217
2-14
038
Mar
supe
lla s
pruc
eipe
rs. c
om. N
orris
200
4
Met
zger
ia c
onju
gata
3
832
310
55-
800
28
Met
zger
ia t
empe
rata
16
71
8-30
515
Moe
rcki
a bl
yttii
1
33
5-14
786
Moe
rcki
a hi
bern
ica
21
115
3-17
475
Myl
ia a
nom
ala
18-
172
4
Myl
ia ta
ylor
ii 1
29
26
5-10
9312
Nar
dia
brei
dler
i 1
1sh
ould
be
chec
ked
agai
n, v
ery
few
r e
cor d
s in
WA
1096
1
Nar
dia
geos
cyph
us
21
213
48-1
539
5
Nar
dia
japo
nica
3
23
1093
-174
07
40
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
36 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Nar
dia
scal
aris
11
91
11
110
-153
925
Odo
ntos
chis
ma
denu
datu
m
12
81
65-
176
9
Pel
lia e
ndiv
iifol
ia
11
822-
1525
4
Pel
lia e
piph
ylla
Pel
lia n
eesi
ana
233
416
41
16
42
5-17
4747
Pla
gioc
hila
asp
leni
oide
s 4
51
1213
11
5-12
7113
Pla
gioc
hila
por
ello
ides
4
198
48
13
32
17
35-
1403
45
Ple
uroc
lada
alb
esce
ns v
ar.
albe
scen
sve
rifie
d 20
051
Por
ella
cor
daea
na
215
151
32
41
132
5-18
0725
Por
ella
nav
icul
aris
3
534
12
124
131
34
31
15-
1807
48
Por
ella
roe
llii
515
11
11
41
93-
707
20
Por
ella
roe
llii f
o. c
rispa
ta
Pre
issi
a qu
adra
ta
270
31
Ptil
idiu
m c
alifo
rnic
um
869
474
98
913
11
54
33
82
185-
1950
103
Ptil
idiu
m c
iliar
e Sp
ecim
en n
eeds
to b
e tra
cked
dow
n,
p rob
ably
P.p
u lc h
e rrim
u mP
tilid
ium
pul
cher
rimum
1
21
Hon
g et
al.
1989
8-12
714
Rad
ula
bola
nder
i 1
722
76
14
28
143
15-
1093
53
Rad
ula
com
plan
ata
939
22
76
23
94
2-12
9235
Rad
ula
obtu
silo
ba s
ubsp
. po
lycl
ada
1Sc
hofie
ld 1
967,
OP
spec
imen
at U
BC87
71
Reb
oulia
hem
isph
aeric
a
Ric
card
ia c
ham
edry
folia
14
51
21
15-
1163
11
Ric
card
ia la
tifro
ns1
131
12
61
11
12
23
5-13
0726
Ric
card
ia m
ultif
ida
133
113
24
12
12
5-86
129
Ric
card
ia p
alm
ata
412
33
41
11
52
55-
1307
42
Ric
cia
beyr
ichi
ana
Susp
ecte
d fro
m sp
ecim
en &
pho
to,
n eed
sver
ifica
ti on
Ric
cia
glau
ca
272
8-15
632
Ric
cioc
arpo
s na
tans
1
791
Sca
pani
a am
eric
ana
536
35
12
12
21
55
15-
1746
44
Sca
pani
a bo
land
eri
26
3898
915
16
45
711
143
712
31
65-
1511
91
Sca
pani
a cu
rta
11
254-
1238
2
Sca
pani
a irr
igua
1
11
1261
-129
92
41
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 37
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Sca
pani
a m
ucro
nata
1
21
12
24-1
325
7
Sca
pani
a ob
scur
a 1
21
need
s ver
ifica
tion
1401
-174
04
Sca
pani
a pa
ludo
sa
24
484
5-17
476
Sca
pani
a su
balp
ina
22
15
211
47-1
747
7
Sca
pani
a ul
igin
osa
12
1348
-134
82
Sca
pani
a um
bros
a 2
35
222
11
12
32
25
5-12
6131
Sca
pani
a un
dula
ta
317
53
14
5-16
2328
Sca
pani
a un
dula
ta v
ar.
oake
sii
11
289-
893
2
Sch
ofie
ldia
mon
ticol
a 4
21
Nor
ris 1
997
1159
-147
87
Trit
omar
ia e
xsec
tifor
mis
2
verif
. R. D
ewey
778
1
Trit
omar
ia q
uinq
uede
ntat
a 4
21
21
verif
. R. D
ewey
822-
1746
6
151
LIV
ERW
OR
T sp
ecie
s fou
nd o
n O
P (2
004)
118
LIV
ERW
OR
T sp
ecie
s on
Rho
ades
list
(199
7)19
5832
1643
2920
2920
1644
930
1528
7980
4775
88
84 L
IVER
WO
RTS
in O
NP
Her
bari
um (O
LYM
) (19
97)
Num
ber o
f LIV
ERW
ORT
spec
ies f
ound
in e
ach
maj
or
vege
tatio
n ty
peN
umbe
r of
LIVE
RWO
RT sp
ecie
s fo
und
per s
ubstr
ate
cate
gory
MO
SSES
Als
ia c
alifo
rnic
a 6
loca
lity
thre
aten
ed43
1
Am
blys
tegi
um s
erpe
ns
11
134-
265
2
Am
phid
ium
cal
iforn
icum
2
147
-104
86
Am
phid
ium
lapp
onic
um
417
9-16
856
Am
phid
ium
mou
geot
ii 4
11
5-12
614
Ana
colia
men
zies
ii 6
147
-115
010
And
reae
a al
pest
ris
11
1163
1
And
reae
a bl
yttii
Sc
hofie
ld 1
967
And
reae
a he
inem
anni
i O
LYM
col
l Nor
ris (v
erify
!)
And
reae
a ni
valis
1
91
1061
-162
07
And
reae
a ru
pest
ris
121
21
27
581-
1807
19
Ano
ecta
ngiu
m a
estiv
um
Scho
field
196
7
Ant
itric
hia
calif
orni
ca
28
13
11
243
-861
12
Ant
itric
hia
curt
ipen
dula
42
82
23
34
61
27
11
38-
1299
57
42
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
38 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Arc
toa
fulv
ella
1
1627
1
Atr
ichu
m s
elw
ynii
191
21
12
21
71
25-
1461
33
Atr
ichu
m u
ndul
atum
2
110
-758
4
Aul
acom
nium
and
rogy
num
3
22
1117
11
31
54
44
0-16
2936
Aul
acom
nium
pal
ustr
e 1
11
62
5-17
4721
Bar
bula
con
volu
ta
11
116
78-1
930
2
Bar
bula
ung
uicu
lata
Sc
hofie
ld 1
967
Bar
tram
ia it
hyph
ylla
2
41
14
194
8-19
308
Bar
tram
ia p
omifo
rmis
5
141
11
21
147
-116
726
Blin
dia
acut
a 1
121
21
145
2-16
7811
Bra
chyd
ontiu
m o
lym
picu
m
Nor
ris 1
997,
WN
HP
Clal
lam
Bra
chyt
heci
um a
lbic
ans
88
31
131
53
11
11
6-19
7833
Bra
chyt
heci
um a
sper
rimum
21
13
11
11
25-
1525
9
Bra
chyt
heci
um b
olan
deri
OLY
M v
erify
(Har
thill
)
Bra
chyt
heci
um c
ollin
um
52
43
1325
-195
09
Bra
chyt
heci
um e
ryth
rorr
hizo
n 1
12
11
111
47-1
950
4
Bra
chyt
heci
um f
rigid
um
59
44
131
32
14
11
31
25-
1746
42
Bra
chyt
heci
um h
olzi
nger
i 1
21
21
31
182
5-16
207
Bra
chyt
heci
um h
ylot
apet
um1
82
13
363
4-16
208
Bra
chyt
heci
um le
iber
gii
54
21
62
19
21
1044
-188
613
Bra
chyt
heci
um o
edip
odiu
m
11
1sin
gle
spec
imen
1511
1
Bra
chyt
heci
um p
lum
osum
Bra
chyt
heci
um r
efle
xum
var
. pa
cific
um2
1325
1
Bra
chyt
heci
um r
ivul
are
31
6-89
76
Bra
chyt
heci
um s
aleb
rosu
m
Bra
chyt
heci
um s
tark
ei
11
165-
1325
2
Bra
chyt
heci
um s
tark
ei v
ar.
expl
anat
um1
861
1
Bra
chyt
heci
um v
elut
inum
4
22
11
11
22
1va
rietie
s not
ent
ered
130-
1950
9
Bra
chyt
heci
um v
elut
inum
var
. ve
lutin
um
Bra
chyt
heci
um v
elut
inum
var
. ve
nust
um2
14
31
1N
orris
199
717
9-18
077
Bro
ther
ella
roe
llii
USF
S, e
xtin
ct?
Pers
. com
. J. H
arpe
l 2 0
04,W
NH
PCl
all .,
Jef f
43
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 39
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Bry
oery
thro
phyl
lum
rec
urvi
rost
re
Bry
um a
mbl
yodo
n
Bry
um a
rgen
teum
1
21
10-
1978
3
Bry
um c
aesp
itici
um
15
1
Bry
um c
alob
ryoi
des
Spen
ce 1
983,
WN
HP
Jeffe
rson
Bry
um c
anar
iens
e N
orris
199
7
Bry
um c
apill
are
86
14
61
42
13
15-
1167
23
Bry
um c
yclo
phyl
lum
1
1348
1
Bry
um d
icho
tom
um
Bry
um f
lacc
idum
Sp
ence
198
3
Bry
um g
emm
ipar
um
11
115
1-15
112
Bry
um li
sae
42
1325
-195
05
Bry
um li
sae
var.
cus
pida
tum
Bry
um m
inia
tum
5
47-1
740
6
Bry
um m
uehl
enbe
ckii
11
5-10
933
Bry
um p
alle
ns
113
481
Bry
um p
alle
scen
s 3
11
170
7-16
295
Bry
um p
seud
otriq
uetr
um
32
12
15-
1745
13
Bry
um t
urbi
natu
m
J. H
arpe
l in
Rhoa
des 1
997
Bry
um w
eige
lii
72
112
13
11
12
76-1
747
26
Bux
baum
ia a
phyl
la
D. V
itt 1
977
data
, in
Rhoa
des 1
997,
W
NH
P Cl
all a
mB
uxba
umia
pip
eri
121
21
11
114
0-13
0710
Bux
baum
ia v
iridi
s Sc
hofie
ld 1
967
Cal
lierg
on c
ordi
foliu
m
51
Cal
lierg
on g
igan
teum
77
81
Cal
lierg
on s
tram
ineu
m
277
8-13
485
Cal
lierg
onel
la c
uspi
data
2
931
Cal
lierg
onel
la li
ndbe
rgii
Nor
ris 1
997
Cam
pyliu
m p
olyg
amum
Cam
pyliu
m s
tella
tum
1
Scho
field
196
713
481
Cam
pylo
pus
intr
ofle
xus
1M
. Hut
ten,
ver
if. J.
Har
pel
1
44
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
40 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Cer
atod
on p
urpu
reus
12
42
22
40-
1978
19
Cirr
iphy
llum
cirr
osum
Cla
opod
ium
bol
ande
ri 5
609
39
81
51
111
1215
123-
1627
52
Cla
opod
ium
cris
pifo
lium
4
1321
53
48
47
15-
1150
40
Cla
opod
ium
whi
pple
anum
1
61
21
13
Scho
field
196
770
7-16
4810
Clim
aciu
m d
endr
oide
s 1
11
5-77
83
Con
osto
mum
tet
rago
num
N
orris
199
7
Cos
cino
don
caly
ptra
tus
Spen
ce 1
983
Cra
tone
uron
fili
cinu
m
34
13
11
347
-162
314
Cru
mia
latif
olia
4
pers
. com
. J. H
arpe
l 200
2, W
NH
P Cl
all a
m0-
472
Cyn
odon
tium
jenn
eri
1Sc
hofie
ld 1
967
977-
1134
2
Cyn
odon
tium
str
umul
osum
N
orris
199
7 sy
n C.
stru
mife
rum
?
Den
droa
lsia
abi
etin
a 3
313
0-45
94
Des
mat
odon
latif
oliu
s 6
24
17
1325
-197
89
Dic
hely
ma
falc
atum
pe
rs. c
om. J
. Har
pel 2
004
Dic
hely
ma
unci
natu
m
Dic
hodo
ntiu
m o
lym
picu
m
24
21
42
1292
-167
87
Dic
hodo
ntiu
m p
ellu
cidu
m
425
21
32
11
25-
1648
33
Dic
rane
lla c
rispa
1
2-28
93
Dic
rane
lla g
revi
llean
a
Dic
rane
lla h
eter
omal
la
203
21
11
12
5-14
3723
Dic
rane
lla h
owei
N
orris
199
7
Dic
rane
lla p
acifi
ca
Scho
field
196
721
11
Dic
rane
lla p
alus
tris
Sc
hofie
ld 1
967
1227
-153
92
Dic
rane
lla r
ufes
cens
1
289-
728
2
Dic
rane
lla s
chre
beria
na
11
Her
man
n 19
6919
01
Dic
rane
lla v
aria
1
1Sc
hofie
ld 1
967
211-
335
2
Dic
rano
wei
sia
cirr
ata
13
22
593
-129
25
Dic
rano
wei
sia
cris
pula
var
. co
nter
min
aN
orris
199
7 (w
e di
d no
t rec
ord
var.)
Dic
rano
wei
sia
cris
pula
var
. cr
ispu
la1
222
21
96
610
44-1
950
20
Dic
ranu
m f
usce
scen
s 8
232
3320
75
56
23
77
33
73
14
5-20
3280
45
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 41
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Dic
ranu
m h
owel
lii
512
1320
231
12
35
96
42
52
20-
2032
60
Dic
ranu
m m
ajus
13
4-16
73
Dic
ranu
m p
allid
iset
um
58
413
34
32
1382
2-17
4624
Dic
ranu
m p
olys
etum
O
LYM
(Har
thill
) det
. K. H
utte
n
Dic
ranu
m r
habd
ocar
pum
Dic
ranu
m s
copa
rium
1
11
83
21
441-
1265
8
Dic
ranu
m t
auric
um
39
153
31
27
37
22
11
10-
1511
37
Dic
ranu
m u
ndul
atum
15
71
Did
ymod
on r
igid
ulus
var
. gr
acili
s
Did
ymod
on t
opha
ceus
Did
ymod
on v
inea
lis2
11
15-
817
8
Did
ymod
on v
inea
lis v
ar.
brac
hyph
yllu
s1
Scho
field
196
76
1
Did
ymod
on v
inea
lis v
ar.
flacc
idus
11
verif
. D. N
orris
199
7 as
D. i
nsul
aris
389
1
Did
ymod
on v
inea
lis v
ar.
vine
alis
122
31
11
21
11
5-11
6018
Dis
celiu
m n
udum
2
verif
. Sch
ofie
ld, W
NH
P Je
ffers
on28
91
Dis
tichi
um c
apill
aceu
m
31
689-
1958
7
Ditr
ichu
m a
mbi
guum
3
11
211-
810
6
Ditr
ichu
m fl
exic
aule
Ditr
ichu
m h
eter
omal
lum
3
18-
1292
2
Ditr
ichu
m m
onta
num
8
34
1Sc
hofie
ld 1
967
822-
1461
5
Ditr
ichu
m p
usill
um
spec
imen
in O
NP
Her
bariu
m, n
ot
verif
ied
Ditr
ichu
m s
chim
peri
11
OLY
M N
orris
WN
HP
Clal
lam
, Gra
ys
Ha r
bor
210
1
Ditr
ichu
m z
onat
um v
ar.
scab
rifol
ium
1ne
eds v
erifi
catio
n16
291
Dre
pano
clad
us a
dunc
us v
ar.
adun
cus
11
8-17
473
Dre
pano
clad
us a
dunc
us v
ar.
knei
ffii
Dre
pano
clad
us s
endt
neri
1N
orris
199
712
711
Dry
ptod
on p
aten
s 1
231
12
61
245
2-17
4623
Enc
alyp
ta a
ffin
is
Scho
field
196
7
Enc
alyp
ta c
iliat
a 2
211-
1150
4
Enc
alyp
ta p
roce
ra
116
301
Enc
alyp
ta r
hapt
ocar
pa
118
071
46
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
42 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Enc
alyp
ta v
ulga
ris
Epi
pter
ygiu
m t
ozer
i 24
12
11
24
14
Nor
ris 1
997
2-12
9229
Euc
ladi
um v
ertic
illat
um
247
1
Eur
hync
hium
ore
ganu
m
128
1417
272
12
16
92
38
31
20-
1629
72
Eur
hync
hium
pra
elon
gum
11
35
417
11
26
31
51
0-11
6033
Eur
hync
hium
pul
chel
lum
var
. pu
lche
llum
1112
116
11
23
114
0-19
5823
Eur
hync
hium
ser
rula
tum
1
verif
icat
ion
need
ed76
71
Fis
side
ns a
dian
thoi
des
257
11
Fis
side
ns b
ryoi
des
31
11
37-7
285
Fis
side
ns c
rispu
s 10
11
51
15-
728
14
Fis
side
ns f
onta
nus
WN
HP
Gra
ys H
arbo
r
Fis
side
ns g
rand
ifron
s 6
2N
orris
199
7, W
NH
P M
ason
104-
822
8
Fis
side
ns p
aupe
rcul
us
WN
HP
Clal
lam
Fis
side
ns v
entr
icos
us
1W
NH
P Cl
alla
m, S
chof
ield
196
7.
Seve
r al r
ecor
dsby
Win
ter e
tal .
2 000
149
1
Fon
tinal
is a
ntip
yret
ica
var.
ant
ipyr
etic
a
Fon
tinal
is a
ntip
yret
ica
var.
ore
gone
nsis
5-12
755
Fon
tinal
is n
eom
exic
ana
11
31
11
5-10
9314
Fon
tinal
is p
atul
aW
alla
ce 1
976
(sus
pect
?)
Fun
aria
hyg
rom
etric
a 2
11
2-25
94
Grim
mia
aff
inis
N
orris
199
7
Grim
mia
ano
don
Spen
ce 1
983
Grim
mia
ano
mal
a 1
Nor
ris 1
997
825
1
Grim
mia
don
nian
a N
orris
199
7
Grim
mia
hol
zing
eri
Her
man
n 19
69
Grim
mia
mon
tana
1
71
9N
orris
199
713
16-1
958
9
Grim
mia
ova
lis
118
071
Grim
mia
pul
vina
ta
Grim
mia
ten
errim
a2
122
612
27-1
950
9
Grim
mia
tor
quat
a 6
266-
1150
9
Grim
mia
tric
hoph
ylla
1
1Sc
hofie
ld 1
967
5-77
53
Gym
nost
omum
rec
urvi
rost
re2
1
47
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 43
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Ham
atoc
aulis
ver
nico
sus
Hed
wig
ia s
tella
taSc
hofie
d 19
67as
H. c
iliat
a
Her
zogi
ella
sel
iger
i 1
11
1Sc
hofie
ld 1
967,
WN
HP
Clal
lam
634-
634
2
Het
eroc
ladi
um d
imor
phum
2
51
140-
1930
6
Het
eroc
ladi
um m
acou
nii
340
41
52
13
15
35-
1307
33
Het
eroc
ladi
um p
rocu
rren
s 5
396
311
61
22
120
87
134-
1627
36
Hom
alia
tric
hom
anoi
des
pers
. com
. J. H
arpe
l 200
4
Hom
alot
heci
um a
eneu
m
41
265-
1082
4
Hom
alot
heci
um a
rena
rium
2
0-47
4
Hom
alot
heci
um f
ulge
scen
s 7
93
12
61
31
15-
1082
27
Hom
alot
heci
um n
evad
ense
5
12
707-
1630
6
Hom
alot
heci
um n
utta
llii
121
34
38
11
15-
707
20
Hom
alot
heci
um p
inna
tifid
um
Hoo
keria
luce
ns
24
61
5-82
217
Hyg
rohy
pnum
bes
tii
11
112
3-16
985
Hyg
rohy
pnum
dur
iusc
ulum
Sc
hofie
ld 1
967
Hyg
rohy
pnum
lurid
um
3W
NH
P Cl
alla
m, J
effe
rson
177-
1348
3
Hyg
rohy
pnum
mol
le
145
2-10
932
Hyg
rohy
pnum
och
race
um
55
22
31
11
5-17
4718
Hyl
ocom
ium
spl
ende
ns
12
59
302
31
21
54
24
73
15-
1347
62
Hyp
num
cal
lichr
oum
Sc
hofie
ld 1
967
Hyp
num
circ
inal
e 4
8838
37
47
93
28
132
511
51
175-
1629
103
Hyp
num
cup
ress
iform
e
Hyp
num
die
ckii
523
29
51
11
12
16
5-16
4843
Hyp
num
lind
berg
ii 2
179-
1623
4
Hyp
num
rev
olut
um
1Sc
hofie
ld 1
967,
OP
spec
imen
at U
BC18
071
Hyp
num
sub
impo
nens
2
319
51
11
23
21
38
12
99-1
048
25
Hyp
num
vau
cher
i
Isop
tery
giop
sis
pulc
hella
2
11
1Sc
hofie
ld 1
967
109-
1630
5
Isot
heci
um c
ardo
tii
112
23
11
1N
orris
199
7. T
his s
peci
es is
not
ac
cept
edby
ITI S
, nor
onth
e B C
list
.13
0-12
9217
Isot
heci
um c
rista
tum
Sc
hofie
ld 1
967
48
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
44 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Isot
heci
um m
yosu
roid
es
321
9823
77
56
115
167
213
22
35-
1624
95
Iwat
suki
ella
leuc
otric
ha
40ve
rif. J
. Har
pel
600-
907
18
Kia
eria
bly
ttii
11
Spen
ce 1
983
1437
1
Kia
eria
fal
cata
1
Scho
field
196
715
091
Kia
eria
sta
rkei
1
163
13
8H
erm
ann
1969
822-
1886
15
Lept
obry
um p
yrifo
rme
11
931
Lept
odic
tyum
rip
ariu
m
1Sc
hofie
ld 1
967
5-42
45
Lesk
ea p
olyc
arpa
2
12
170
7-16
202
Lesk
eella
ner
vosa
Sc
hofie
ld 1
967
Leuc
olep
is a
cant
hone
uron
5
28
611
21
36
51
23
11
5-98
743
Mee
sia
ulig
inos
a W
NH
P Cl
alla
m, J
effe
rson
Met
anec
kera
men
zies
ii 3
261
14
61
14
493
-948
25
Mni
um a
mbi
guum
1
189
7-18
072
Mni
um b
lytt
ii 1
11
Scho
field
196
712
71-1
630
2
Mni
um ly
copo
dioi
des
31
21
2Sc
hofie
ld 1
967
103-
1150
5
Mni
um m
argi
natu
m
11
179-
689
3
Mni
um s
pinu
losu
m
55
27
214
15
11
12
538
9-16
2930
Mni
um t
hom
soni
i1
11
Scho
field
196
7 as
M. o
rthor
rhyn
chum
1048
-180
73
Myu
rella
jula
cea
Scho
field
196
7, W
NH
P Je
ffers
on,
Mas
onN
ecke
ra d
ougl
asii
3811
25
715
25
93
15-
845
37
Olig
otric
hum
alig
erum
21
11
25-
1391
15
Olig
otric
hum
her
cyni
cum
1
11
685-
822
2
Olig
otric
hum
par
alle
lum
4
14
12
128
9-15
5113
Onc
opho
rus
vire
ns
Scho
field
196
7
Onc
opho
rus
wah
lenb
ergi
i N
orris
199
7
Ort
hoth
eciu
m c
hrys
eum
W
NH
P Cl
alla
m
Ort
hotr
ichu
m a
ffin
e 3
12
unve
rif. O
LYM
Bro
wn,
Har
thill
758-
1233
3
Ort
hotr
ichu
m a
lpes
tre
415
3-19
584
Ort
hotr
ichu
m a
nom
alum
1
1958
1
Ort
hotr
ichu
m c
onsi
mile
22
31
75
31
22-
1044
23
Ort
hotr
ichu
m la
evig
atum
3
1630
-180
72
49
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 45
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Ort
hotr
ichu
m ly
ellii
1
351
73
13
41
22-
1271
31
Ort
hotr
ichu
m o
btus
ifoliu
m
143
-778
4
Ort
hotr
ichu
m p
ellu
cidu
m
31
778-
1807
3
Ort
hotr
ichu
m p
ulch
ellu
m
115
11
22
11
13
15-
1271
16
Ort
hotr
ichu
m p
ylai
sii
WN
HP
Clal
lam
Ort
hotr
ichu
m r
ivul
are
2N
orris
199
726
-897
4
Ort
hotr
ichu
m r
upes
tre
126
5-11
502
Ort
hotr
ichu
m s
peci
osum
1
113
01
Ort
hotr
ichu
m s
tria
tum
1
218
1
Pal
ustr
iella
com
mut
ata
11
1Sc
hofie
ld 1
967,
Nor
ris 1
997
as
Crat
oneu
r on
com
mut
atum
var .
211-
1623
5
Pal
ustr
iella
fal
cata
Nor
ris 1
997
as C
rato
neur
on
com
mut
atum
var .
f al c
atum
Par
aleu
cobr
yum
ene
rve
Phi
lono
tis c
apill
aris
72
81
Phi
lono
tis f
onta
na v
ar.
amer
ican
a1
42
1N
orris
199
726
5-16
9810
Phi
lono
tis f
onta
na v
ar.
caes
pito
sa3
12
11
822-
1747
8
Phi
lono
tis f
onta
na v
ar.
font
ana
84
14
11
123-
1747
18
Phi
lono
tis f
onta
na v
ar.
pum
ila1
1N
orris
199
711
64-1
685
4
Phi
lono
tis m
uehl
enbe
rgii
118
861
Phi
lono
tis y
ezoa
na
Nor
ris 1
997
822-
822
2
Phy
scom
itriu
m p
yrifo
rme
Rhoa
des 1
994
Pla
giob
ryum
zie
riiSc
hofie
ld 1
967
Pla
giom
nium
cili
are
11
1N
orris
199
713
4-16
52
Pla
giom
nium
elli
ptic
um
189
71
Pla
giom
nium
insi
gne
95
12
213
12
43
55
25
15-
1271
44
Pla
giom
nium
med
ium
1
41
1Sc
hofie
ld 1
967
5-12
996
Pla
giom
nium
ros
trat
um
25
12
21
12
159-
1164
17
Pla
giom
nium
ven
ustu
m
21
162
14
51
22
326
-822
24
Pla
giop
us o
eder
iana
Sc
hofie
ld 1
967
1
Pla
giot
heci
um c
avifo
lium
3
15
13
1N
orris
199
75-
1627
10
Pla
giot
heci
um d
entic
ulat
um
23
36
32
32
5-16
2014
Pla
giot
heci
um la
etum
7
622
2620
54
59
24
91
61
28
5-16
7854
50
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
46 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Pla
giot
heci
um p
ilife
rum
7
71
11
11
299
-156
117
Pla
giot
heci
um u
ndul
atum
4
1827
133
41
15
65
11
72
13
5-12
7159
Pla
tydi
ctya
jung
erm
anni
oide
s 1
31
147
-163
09
Pla
tyhy
pnid
ium
rip
ario
ides
2
WN
HP
Gra
ys H
arbo
r1
Ple
urid
ium
sub
ulat
um
Nor
ris 1
997
Ple
uroz
ium
sch
rebe
ri 3
21
16
18-
1238
17
Pog
onat
um c
onto
rtum
17
12
16
22
12
21
5-82
227
Pog
onat
um u
rnig
erum
6
41
21
15-
1740
17
Poh
lia a
nnot
ina
31
117
9-15
114
Poh
lia a
trop
urpu
rea
41
OLY
M c
oll.
Har
thill
5-14
763
Poh
lia b
olan
deri
134
12
24
41
289-
1978
17
Poh
lia b
olan
deri
var.
ser
iata
11
1511
-195
82
Poh
lia c
ampt
otra
chel
a1
1ne
eds v
erif.
1511
1
Poh
lia c
olum
bica
1
need
s ver
if.10
31
Poh
lia c
ruda
14
103
41
33
48-
1807
26
Poh
lia d
rum
mon
dii
Poh
lia e
long
ata
OLY
M c
oll.
(Har
thill
)
Poh
lia e
rect
a 1
1N
orris
199
716
98-1
746
2
Poh
lia f
ilum
1
6-13
482
Poh
lia le
scur
iana
O
LYM
col
l. H
arth
ill
Poh
lia lo
ngib
ract
eata
10
18-
512
7
Poh
lia lu
dwig
ii 1
1N
orris
199
714
761
Poh
lia n
utan
s 10
12
59
24
82
11
12
5-16
7820
Poh
lia o
btus
ifolia
1
111
631
Poh
lia p
acifi
ca
51
8-51
24
Poh
lia p
rolig
era
11
1147
1
Poh
lia v
exan
s Sc
hofie
ld 1
967
Poh
lia w
ahle
nber
gii
272
11
12
11
15-
1678
23
Pol
ytric
hast
rum
alp
inum
var
. al
pinu
m13
101
39
22
61
21
14
21
25-
1629
39
Pol
ytric
hum
com
mun
e 2
11
81
5-13
4814
Pol
ytric
hum
for
mos
um
134-
182
3
51
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 47
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Pol
ytric
hum
juni
perin
um
122
13
61
13
21
20-
1740
26
Pol
ytric
hum
long
iset
um
36
21
11
1ve
rify
spec
imen
s5-
1698
9
Pol
ytric
hum
lyal
lii
43
48
182
5-19
508
Pol
ytric
hum
pili
feru
m
207
112
810
121
4-20
3229
Pol
ytric
hum
sex
angu
lare
Sc
hofie
ld 1
967
Pol
ytric
hum
str
ictu
m
Nor
ris 1
997,
WN
HP
Clal
lam
99-1
572
Por
otric
hum
big
elov
ii 17
32
14
34
5-10
4817
Por
otric
hum
van
couv
erie
nse
21
134-
165
2
Pse
udob
raun
ia c
alifo
rnic
a Sc
hofie
ld 1
967
Pse
udol
eske
a at
richa
5
11
11
799-
1509
6
Pse
udol
eske
a ba
ileyi
1
31
484
5-12
383
Pse
udol
eske
a in
curv
ata
12
11
11
1261
-162
74
Pse
udol
eske
a in
curv
ata
(inte
rmed
iate
for
m)
81
17
799-
1678
8
Pse
udol
eske
a in
curv
ata
var.
gig
ante
a4
11
1082
-162
04
Pse
udol
eske
a in
curv
ata
var.
incu
rvat
a1
61
11
31
1002
-195
88
Pse
udol
eske
a pa
tens
2
142
13
19
31
165-
1747
15
Pse
udol
eske
a ra
dico
sa
11
11
1147
-143
72
Pse
udol
eske
a ra
dico
sa v
ar.
radi
cosa
15
11
13
110
44-1
950
7
Pse
udol
eske
a sa
vian
a 1
1509
1
Pse
udol
eske
a st
enop
hylla
3
171
14
54
558-
1886
20
Pse
udos
cler
opod
ium
pur
um
M. H
utte
n da
ta
Pse
udot
axip
hyllu
m e
lega
ns
2623
824
1011
15
21
35
22
55
27
5-17
4567
Pte
rigyn
andr
um f
ilifo
rme
16
51
182
2-19
509
Pte
rogo
nium
gra
cile
Rac
omitr
ium
aci
cula
re
181
11
10-1
391
18
Rac
omitr
ium
aff
ine
51
194
8-12
335
Rac
omitr
ium
aqu
atic
um
201
verif
. J. H
arpe
l, W
NH
P Cl
alla
m21
2-96
617
Rac
omitr
ium
bre
vipe
s 1
244
33
51
21
257
4-18
8625
Rac
omitr
ium
can
esce
ns v
ar.
cane
scen
s
Rac
omitr
ium
elo
ngat
um
1113
113
11
44
13
0-16
2933
Rac
omitr
ium
eric
oide
s 3
21
14
176-
1539
7
52
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
48 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Rac
omitr
ium
fas
cicu
lare
Rac
omitr
ium
het
eros
tichu
m
124
12
22
12
115
1-16
2428
Rac
omitr
ium
lanu
gino
sum
6
539-
1347
4
Rac
omitr
ium
mac
ouni
i 11
12
11
141
9-17
4515
Rac
omitr
ium
mic
roca
rpon
2
101
21
31
need
s ver
ifica
tion
767-
1886
11
Rac
omitr
ium
obe
sum
2
11
Nor
ris 1
997
1325
-162
74
Rac
omitr
ium
occ
iden
tale
19
42
1282
2-16
7813
Rac
omitr
ium
pac
ificu
m
1ne
eds v
erifi
catio
n10
931
Rac
omitr
ium
pyg
mae
um
Nor
ris 1
997
Rac
omitr
ium
sud
etic
um
21
211
63-1
678
5
Rac
omitr
ium
var
ium
3
141
32
12
37-1
978
23
Rha
bdow
eisi
a cr
ispa
ta
OLY
M c
oll.
Har
thill
(miss
ing?
) un
likel
yac
cSc
hofie
ld20
04R
hizo
mni
um g
labr
esce
ns
54
1652
45
52
23
137
12
81
21
5-12
7169
Rhi
zom
nium
mag
nifo
lium
5
11
82
21
21
12
5-16
9833
Rhi
zom
nium
nud
um
41
11
155
13
11
3ve
rif. J
. Har
pel
202-
1698
26
Rhi
zom
nium
pse
udop
unct
atum
1
18-
1623
2
Rhi
zom
nium
pun
ctat
um
Nor
ris 1
997
Rhy
tidia
delp
hus
lore
us
44
1516
235
73
23
54
22
61
85-
1476
67
Rhy
tidia
delp
hus
squa
rros
us
31
310
31
23
21
493
-123
321
Rhy
tidia
delp
hus
subp
inna
tus
21
11
5-11
475
Rhy
tidia
delp
hus
triq
uetr
us
42
75
192
34
32
26
12
0-11
6743
Rhy
tidio
psis
rob
usta
2
81
727
74
413
21
19
441-
1678
52
Rhy
tidiu
m r
ugos
um1
Spen
ce 1
986
(nea
rly e
xtin
ct?)
, WN
HP
Clal
l am
1807
1
Roe
llia
roel
lii
63
11
204
41
111
17
758-
1950
35
San
ioni
a un
cina
ta
33
13
13
897-
1698
9
Sar
men
thyp
num
sar
men
tosu
m
Sch
istid
ium
aga
ssiz
ii 1
Nor
ris 1
997
151
1
Sch
istid
ium
apo
carp
um
115
13
118
0-19
5017
Sch
istid
ium
mar
itim
um
15
0-26
4
Sch
istid
ium
riv
ular
e va
r. la
tifol
ium
Sch
istid
ium
riv
ular
e va
r. r
ivul
are
17
126
-129
99
53
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 49
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Sch
istid
ium
ten
erum
Sch
isto
steg
a pe
nnat
a 10
1ve
rif. J
. Har
pel,
WN
HP
Gra
ys H
arbo
r, Je
ff er s
on17
6-25
57
Scl
erop
odiu
m c
espi
tans
var
. ce
spita
ns1
12-
153
3
Scl
erop
odiu
m c
espi
tans
var
. su
blae
ve
Scl
erop
odiu
m o
btus
ifoliu
m
215
11
12
5-17
4719
Scl
erop
odiu
m t
oure
tii v
ar.
colp
ophy
llum
Nor
ris 1
997
Scl
erop
odiu
m t
oure
tii v
ar.
tour
etii
2Sc
hofie
ld 1
967
26-1
167
5
Sco
uler
ia a
quat
ica
111
247
-156
316
Sph
agnu
m a
lask
ense
And
rus &
Jans
sens
200
3,
OP
spec
imen
atU
B CS
phag
num
ang
ustif
oliu
m(s
yn. S
. rec
urvu
m C
rum
198
4)
Sph
agnu
m a
ustin
ii W
NH
P Cl
alla
m, O
P sp
ecim
en a
t UBC
Sph
agnu
m c
apill
ifoliu
m
1N
orris
199
78-
134
3
Sph
agnu
m c
apill
ifoliu
m v
ar.
tene
llum
18-
172
4
Sph
agnu
m f
alla
x N
orris
199
7
Sph
agnu
m f
imbr
iatu
m
153
1
Sph
agnu
m f
uscu
m
OP
spec
imen
at U
BC15
3-15
72
Sph
agnu
m g
irgen
sohn
ii 2
81
15-
1079
13
Sph
agnu
m h
enry
ense
Sph
agnu
m m
agel
lani
cum
8-
162
Sph
agnu
m m
endo
cinu
m
OP
spec
imen
at U
BC8-
778
3
Sph
agnu
m p
acifi
cum
Fl
atbe
rg 1
989
OP
spec
imen
at U
BC
(in
cl.S
chof
i el d
196
7 r e
cor d
ofS.
Sph
agnu
m p
alus
tre
13
35-
424
8
Sph
agnu
m p
apill
osum
1
18-
172
5
Sph
agnu
m q
uinq
uefa
rium
1
unve
rif. s
pec.
OLY
M H
arth
ill17
21
Sph
agnu
m r
usso
wii
153
1
Sph
agnu
m s
quar
rosu
m
11
51
15-
1561
17
Sph
agnu
m s
ubse
cund
umN
orris
199
777
81
Sph
agnu
m s
ubse
cund
um v
ar.
rufe
scen
s1
25-
778
6
Sph
agnu
m t
eres
2
1227
-122
72
Sph
agnu
m w
arns
torf
ii
Spl
achn
um a
mpu
llace
um
Nor
ris 1
997
157-
176
2
54
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
50 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Spl
achn
um s
phae
ricum
15
7-28
93
Spl
achn
um v
ascu
losu
m
Tay
loria
ser
rata
1
21
12
11
1N
orris
199
7, W
NH
P Cl
alla
m5-
1620
7
Tet
raph
is g
enic
ulat
a 14
13
verif
. J. H
arpe
l, W
NH
P G
rays
Har
bor,
Jeffe
rso n
85-5
6515
Tet
raph
is p
ellu
cida
43
21
512
46
15-
1164
36
Tet
rapl
odon
ang
usta
tus
need
s ver
ifica
tion
157
1
Tet
rapl
odon
mni
oide
s 1
12
Nor
ris 1
997,
WN
HP
Clal
lam
5-16
76
Tha
mno
bryu
m n
ecke
roid
es
32
1N
orris
199
717
9-82
23
Tim
mia
aus
tria
ca
43
11
21
310
1-16
2018
Tim
mia
meg
apol
itana
Nor
ris 1
997
Tim
mie
lla c
rass
iner
vis
47-7
282
Tor
tella
fra
gilis
Tor
tella
tor
tuos
a 6
182
2-19
586
Tor
tula
am
plex
a N
orris
199
7 (a
s S.a
mpl
exa)
Tor
tula
bol
ande
ri 1
51
Tor
tula
mur
alis
5
11
0-19
784
Tor
tula
nor
vegi
ca
21
13
112
33-1
886
5
Tor
tula
pap
illos
issi
ma
21
22
110
82-1
958
5
Tor
tula
prin
ceps
1
1Sc
hofie
ld 1
967
2-26
54
Tor
tula
rur
alis
4
74
13
32
0-19
7820
Tor
tula
sub
ulat
a N
orris
199
7
Tra
chyb
ryum
meg
aptil
um
21
41
22
26-
1167
16
Tric
hodo
n cy
lindr
icus
4
11
11
93-1
271
4
Trip
tero
clad
ium
leuc
ocla
dulu
m
Nor
ris 1
997
Ulo
ta m
egal
ospo
ra
106
37
25
34
21
13
63
123-
1461
72
Ulo
ta o
btus
iusc
ula
432
75
21
16
12-
931
40
Ulo
ta p
hylla
ntha
2
154-
268
War
nsto
rfia
exa
nnul
ata
13
Scho
field
196
75-
1539
9
War
nsto
rfia
flu
itans
2
15-
1275
8
Wei
ssia
con
trov
ersa
1
265-
1159
3
Zyg
odon
viri
diss
imus
55
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
Results 51
Scie
ntifi
c na
me
Bare SoilRockEpiphyte
Wood Debris Fine Organic
ABAMABAM dry
ABAM wet
ABLA
ALRUBOG
Parkland
PISI
PSMEPSME dry
PSME wet
TSHETSHE dry
TSHE wet
TSME
Maj
or v
eget
atio
n ty
pe
Subs
trat
e ca
tego
ry
OP localities
OP
elev
atio
n ra
nge
(met
ers)
Sour
ce/c
omm
ent
Rhoades OLYM
(FH
M- s
tyle
sam
plin
g pr
otoc
ol)
1
2
5
3
4
67
Zyg
odon
viri
diss
imus
var
. ru
pest
ris3
179-
179
3
Zyg
odon
viri
diss
imus
var
. vi
ridis
sim
us
419
MO
SS sp
ecie
s fou
nd o
n O
P (2
004)
322
MO
SS sp
ecie
s on
Rho
ades
list
(199
7)59
8310
333
8850
5063
4031
8933
4449
2917
119
588
9913
5
262
MO
SSES
in O
NP
Her
bari
um (O
LYM
) (19
97)
Num
ber o
f MO
SS sp
ecie
s fou
nd in
eac
h m
ajor
veg
etat
ion
type
Num
ber o
f MO
SS
spec
ies f
ound
per
su
bstra
te c
ateg
ory
146
220
212
8620
613
514
315
611
996
217
8211
712
189
320
449
352
292
338
Tota
l num
ber o
f lic
hen
and
bryo
phyt
e sp
ecie
s fou
nd in
eac
h m
ajor
veg
etat
ion
type
Tota
l num
ber o
f lic
hens
and
bry
ophy
tes
foun
d pe
r sub
strat
e ca
tego
ry12
83 T
otal
# sp
ecie
s doc
umen
ted
from
the
Oly
mpi
c Pe
nins
ula
(200
4)
905
Lic
hen
and
Bryo
phyt
e Sp
ecie
s on
Rho
ades
OP
list (
1997
)
498
Lic
hen
and
Bryo
phyt
e Sp
ecie
s in
ON
P H
erba
rium
(OLY
M)(1
997)
56
Tabl
e 1.
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 5
2.)
1 Che
ckm
ark
indi
cate
s spe
cies
repo
rted
by R
hoad
es (w
ritte
n co
mm
., 19
97)
2 Spe
cies
with
out d
ata
in o
ther
col
umns
but
with
a c
heck
mar
k un
der O
LYM
ar
e in
the
ON
P co
llect
ion,
pre
-dat
ing
this
pro
ject
. Spe
cies
with
dat
a in
oth
er
colu
mns
are
par
t of t
his p
roje
ct.
3 Spe
cies
als
o co
llect
ed b
y ot
hers
are
cite
d; li
chen
s with
“O
P sp
ecim
en
depo
site
d in
her
bariu
m B
G”
wer
e co
llect
ed b
y T.
Tøn
sber
g an
d ar
e in
the
Uni
vers
ity o
f Ber
gen’
s col
lect
ion.
4 Ele
vatio
n ra
nge
data
are
bas
ed o
n re
cord
s gen
erat
ed d
urin
g th
is p
roje
ct.
Elev
atio
n re
cord
s in
the
NPS
cur
ator
ial d
atab
ase
wer
e ex
clud
ed b
ecau
se o
f in
adeq
uaci
es in
the
spat
ial d
ata.
5 The
num
ber o
f col
lect
ion
loca
litie
s ind
icat
es u
niqu
e si
tes,
alth
ough
uni
que
site
s may
be
clos
e to
geth
er.
6 Tal
lies a
re b
ased
on
FHM
-sty
le sa
mpl
ing
prot
ocol
; veg
etat
ion
type
s are
bo
g (B
OG
), re
d al
der (
ALR
U),
Sitk
a sp
ruce
(PIS
I), w
este
rn h
emlo
ck (T
SHE)
, D
ougl
as-f
ir (P
SME)
, silv
er fi
r (A
BA
M),
mou
ntai
n he
mlo
ck (T
SME)
, and
a
varie
ty o
f tim
berli
ne h
abita
ts ra
ngin
g fr
om sc
ree
slop
es to
alp
ine
tund
ra
(Par
klan
d).
7 Gen
eral
ized
subs
trate
s wer
e pr
ovid
ed to
giv
e an
app
roxi
mat
e id
ea o
f the
sp
ecie
s’ m
icro
habi
tat.
52 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Other products
Several products resulted from this project in addition to a significantly enhanced list and herbarium of cryptogam species for ONP.
• Field GuideA field guide entitled “101 Common Mosses, Liverworts,
and Lichens of the Olympic Peninsula” (Hutten and others, 2001) is a pocket-sized, waterproof guide to cryptogams of the Olympic Peninsula, and it is also useful throughout western Washington. It is easy to use, with close-up color photos and non-technical descriptions that offer tips for identification. Species are grouped by substrate, and elevational range is indicated for each. It includes a glossary and diagrams to teach new terms, and differentiates 140 species. This field guide is for sale to the public and will also be used to train vegetation-monitoring crews in the future.
• Illegal Commercial Moss HarvestsImages and reports describing two significant incidents
of illegal harvest of cryptogams from the Park are available to ONP staff members at the park-accessible network address I:\all\crypto\Commercial Moss Harvest\ (restricted access).
• Fact SheetA four-page fact sheet describing cryptogams, their eco-
logical role, and their need for conservation is available from USGS as a PDF file at http://fresc.usgs.gov/products/fs/fs-154-02.pdf, November 2005. This fact sheet is also available in hard copy from USGS-FRESC, 777 NW 9th St., Suite 400, Corvallis, OR 97330.
Images that have been provided to other agencies can be seen at http://www.or.blm.gov/surveyandmanage/about.htm, temporarily unavailable November 2005 and http://www.fs.fed.us/pnw/pubs/gtr573.pdf, November 2005. Other images have been requested for use in a general biology textbook, and 50 images have been published (Hutten, 2003; http://www.cnps.org/publications/fremontia/Fremontia_Vol31-No3.pdf, November 2005).
• WebsitesGeneral project information is available at http://fresc.
usgs.gov/olympic/research/indivproj.asp?OFSProj_ID=12, November 2005. Results from the ONP non-vascular plant inventory through 2001 can be found at http://ocid.nacse.org/nbii/crypto/, November 2005. This is an online database searchable by watershed (to generate species lists), genus, species (to generate locality listings), and so forth. Several hundred low-resolution photographs can be viewed and down-loaded from this site (higher resolution cryptogam images are accessible to ONP employees at the ONP network drive address I:\all\crypto\images, restricted access).
• GIS LayerAll records of cryptogam species and their locations are
available as a data layer in the ONP GIS system.
• MetadataMetadata for this project and the resulting database can
be found at http://mercury.ornl.gov/nbii/full.jsp?index=0&recidx=0&pageMax=10, November 2005.
• Collaboration with Other AgenciesData of federally listed Survey and Manage species
found in ONP have been shared with the Northwest Forest Plan Interagency Bryologist and Lichen Coordinator with per-mission of the ONP Chief of Natural Resource Management. Species included in the Survey and Manage List required special surveys in areas on USFS and BLM land where ground-disturbing activities are planned. Sharing data con-tributed to the understanding of the habitat requirements and distribution of these species, which is used to refine search protocols. ONP data were included in species habitat models, which predict likely sites for rare species (R. Lesher, written commun., 2002). Some species (such as Leptogium brebis-sonii, Ptilidium californicum, and Ulota megalospora) were dropped from the former Survey and Manage List because they were either common or because habitat data showed that they were not exclusively associated with old-growth forests, a requirement for the federal listing associated with the North-west Forest Plan. Some former Survey and Manage species are locally common in ONP and rare elsewhere in the PNW (for instance, Schistostega pennata, Tetraphis geniculata, Diplophyllum plicatum [fig. 4], Kurzia spp., and Herbertus aduncus [fig. 5]). Botanists of several agencies have visited populations of former Survey and Manage species in ONP to improve their ability to locate these species elsewhere in the region. Washington State Department of Natural Resources, USFS (http://www.fs.fed.us/pnw/pubs/gtr573.pdf, November 2005), and BLM (http://www.or.blm.gov/surveyandmanage/about.htm, temporarily unavailable November 2005) have also used images generated during this project.
Some rare species found in ONP were new to Washing-ton and have been proposed and subsequently added to the Washington Natural Heritage Program (WNHP) list. Thus far, the authors have proposed the lichens Bunodophoron melano-carpum (fig. 6), Spilonema sp.1 (Goward, 1999), and Pseu-docyphellaria mallota (fig. 7). Suggested for listing review is Bryoria tenuis, a very rare species in Washington (possibly new to Washington), with only one or two sites on the OP. Proposed mosses include several species in the Splachna-ceae, including Splachnum ampullaceum (fig. 8). To date, the WNHP list has not included liverworts, but a working list is currently being compiled and ONP data can be made available to support this effort.
ONP has made its data available to other agencies, but informal data requests by ONP are not always answered.
Results 53
Figure 5. Herbertus aduncus.
Figure 6. Bundophoron melanocarpum.
Figure 7. Pseudocyphellaria mallota.
Figure 8. Splachnum ampullaceum.
Figure 4. Diplophyllum plicatum.
54 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
As a result, ONP has no information on the location of the following critically endangered species: Brachydontium olympicum (WNHP G3S1; see Appendix A for explanation of species status), Dermatocarpon luridum (WNHP G?S2), and Umbillicaria scholanderi (WNHP G1S1).
Discussion
Cryptogam Diversity of the Olympic Peninsula
Because of its tremendous habitat complexity, pristine condition, and moderate maritime climate, ONP has an aston-ishing diversity of non-vascular plants. In the Hoh rainforest, a single old-growth Sitka spruce may support more than 60 bryophyte and lichen species and a few epiphytic vascular plants (Hutten, unpub. data, 2001). More than 1,280 spe-cies of mosses, liverworts, hornworts, and lichens have been catalogued from the OP (Rhoades, 1997, supplemented with data from this project), and species new to the area are still frequently discovered. Within ONP, collections and locality data are available for some 900 species. The sheer biomass of bryophytes and lichens is as impressive as the species diver-sity. ONP is internationally known for the “Hall of Mosses”, one of few places in the world where one cannot see the forest for the moss carpets.
It is not just the lush rainforests that support a high abundance and diversity of cryptogams. The unique bioge-ography of the OP includes a variety of other habitats with large numbers of species, including many rare and endemic vascular plants, relative to its geographic area (Houston and others, 1994; Buckingham and others, 1995). In fact, the OP has the greatest number of rare vascular plant taxa listed by WNHP as compared to any geographic area within Washing-ton (http://www.nps.gov/olym/ea/Fmp2/fmpaf2.htm, November 2005). The number of rare non-vascular plants is even higher than vascular plants, and can also be attributed to the broad range of habitats on the OP. Intact examples of many of these habitats are rare elsewhere due to human activity.
Bryophytes and lichens have a high degree of habitat and substrate specificity, which also contributes to the high diversity and frequency of rare taxa on the OP. Though some species are clearly generalists, many bryophytes and lichens are decidedly microhabitat specific. For example, Splachnum ampullaceum and S. sphaericum are mosses found only on excrement in open and perennially moist Sphagnum-domi-nated coastal bogs. Both species were proposed for listing in WNHP, as there are few, if any, other known occurrences in Washington. Occurrences of Tetraplodon mniodes (G5 S1) and Tayloria serrata (G4? S1; fig. 9) on the OP are primarily on moist and exposed coyote excrement. The widespread crustose lichen Fellhanera bouteillei is one of the few epiphylls found in this area and lives on live needles of conifers in moist for-ests. With species occupying such small ecological niches, the wide variety of habitats on the OP could indeed be expected to harbor a multitude of cryptogams.
Rare Species on the Olympic Peninsula
Determining whether a given species is common, infre-quent, or rare requires a large amount of distributional data. Despite having access to more than 20,000 data points for non-vascular plants on the OP, the data are still inadequate for determining the status of a flora of more than 1,250 species, forcing reliance on data from elsewhere in the region. The most relevant source is the list of endangered, threatened, and sensitive plants in Washington, produced by the Washington Natural Heritage Program (WNHP) (1997), a Washington State Department of Natural Resources project. Documents associated with the former Survey and Manage program of the Northwest Forest Plan provide information on species thought to be rare due to their association with old-growth forests. The authors also included the British Columbia (BC) listing of red- and blue-listed mosses and lichens. The BC status data are particularly relevant for those species at the northern extent of their range in this area.
The current list of rare species reported from the OP (Appendix A) includes data from this project, supplemented with select literature reports and personal communications with other lichenologists and bryologists. More than 220 bryo-phytes and lichens found on the ONP are listed as rare in the Washington and Oregon Heritage Plans (ONHIC, 2004) and in former Survey and Manage documents (about 18 percent of the OP flora). The authors expect that the total number of rare species will increase in the future for a number of reasons.
1. There is currently no status information for Wash-ington liverwort species, despite a recent catalog (Hong, 2002). Liverworts, therefore, have not yet been incorporated into the WNHP. The authors used data from ONHIC as an indicator of liverwort rarity (Appendix A), but these Oregon data cannot be used for Washington without caution. There are several OP species that do not occur in Oregon (such as Odontos-chisma denudatum and Blepharastoma arachnoideum [fig. 10]). For lack of Washington distribution data, such species were omitted from our list, although they almost certainly belong on it.
2. Some species are so poorly known that their distribu-tional status cannot yet be determined. Such species are omitted from this list of rare species (Appendix A). For example, Stigonema sp. 1, (Goward, 1999) prob-ably is genuinely rare, but lichenologists have not been aware of this species for very long.
3. The ONP/OP inventory is far from complete, and many crustose lichens and less common bryophytes and macro-lichen species are yet to be recorded.
There are also species on the WNHP list that appear to be quite frequent on the OP (such as the lichens Ahtiana pal-lidula and Usnea wirthii and the liverworts Douinia ovata and Diplophyllum albicans). The status of these lichens may be
Discussion 55
updated if evidence shows that they are also frequent else-where in Washington. Both liverworts were removed recently from the former Survey and Manage list. The moss Racomi-tium aquaticum is now considered to be the distinct species Racomitrium ryszardii, an endemic to the PNW found mainly on the western part of the OP (Bednarek-Ochyra, 2000). The lichen Karnefeltia californica is apparently over-reported in Washington because of mis-identifications; there are perhaps only four or five accepted locations in Washington (L. Geiser, written commun., 2004), hence its status should be G2S1.
Surveyor attention often focuses on species listed as rare. The Survey and Manage species in particular receive much more attention than other species and are specifically sought in the field. On the OP, for example, the authors have collected the rare lichen Pseudocyphellaria rainierensis (fig. 11) from more locations than the common P. anthraspis. It is therefore necessary to use caution in drawing conclusions from the collection frequency of these species on the OP, as indicated in Appendix A. However, a few species do appear to be more frequent here than elsewhere in Washington and Oregon, and future work may demonstrate that they are not truly rare on the OP (for example, Metzgeria temperata and Platismatia lacunosa).
Endemic Species
Clues from biogeography help us to understand patterns of endemism. Endemic species are those with narrow distribu-tional ranges, and are considered special for being unique to a region. Endemism may occur when an area has been isolated for a significant time or serves as a refuge for relict popula-tions of a formerly common species following widespread extinction. Vascular plants endemic to OP were thought to have been isolated by its island-like geography and its poten-tial to have been a refuge during the last ice age (Buckingham and others, 1995). Whether the distribution of non-vascular endemic plants supports this hypothesis remains to be seen.
Among the non-vascular flora, approximately 105 moss species and varieties are endemic to western North America, representing about 18 percent of the moss flora (Lawton, 1971; Schofield, 1985). Of these, some 20 mosses are found west but not east of the Cascade Range (Lawton, 1971). The proportion of endemic liverworts is similar, about 16 percent (Schofield, 1985). Nine moss and three (monotypic) liver-wort genera are endemic to Pacific North America (Scho-field ,1985; 2002). The proportion of the lichen flora that is endemic to western North America is high, but the number is unknown to the authors. At present, no mosses, liverworts, or lichens are known to be endemic exclusively to the Olympic Mountains. In Pacific North America, almost all endemic bryophytes occupy geographic areas that are larger than the OP, hence the Peninsula may not be sufficiently isolated to produce endemic bryophytes. With respect to lichen ende-mism on the OP, Rhoades (1997) comments that “the general response from lichenologists concerning endemism is that the
Olympic Peninsula has a diverse and abundant lichen flora but that, with perhaps a few exceptions, the flora is no different from that of nearby areas.”
Even though there is no current evidence of endemism in the OP non-vascular flora, it may be too early to draw a firm conclusion. Many of the areas suspected of having been glacial refugia in ONP have not been surveyed. The state of knowledge of cryptogam distribution is also still dynamic, as new species are frequently added to the non-vascular plant lists of the OP, Washington, and North America. Species altogether new to science are described frequently, particularly among the crustose lichens. Often, with the increased attention brought on by the publication of a new species description, the species is soon reported from other areas. For example, the moss Brachydontium olympicum was initially believed to be restricted to the Olympics when discovered by Dr. Frye in 1907. It was later found in Oregon, Washington, British Columbia, and Japan. The small fruticose lichen Spilone-mella americana was recently described and placed in a new genus. Originally collected adjacent to the Hoh River (type locality), it is now also known from other areas in Pacific North America (Henssen and Tønsberg, 2000). The ‘arboreal hairball,’ another fruticose lichen species, tentatively called Stigonema sp. 1, was included in a species key for British Columbia by Goward (1999). During this project it was found on Sitka spruce branches from the upper canopy in the Hoh and Quinault rainforests, and it was also found in Oregon (B. McCune,written commun., 2002). During this project, two macro-lichens in the genus Dermatocarpon were found in ONP that are thought to be undescribed. One liverwort in the genus Scapania did not fit any key we had available. If these are undescribed, these species may be endemics, or they may be found elsewhere now that taxonomists are aware of them.
Conservation of Cryptogams
• Regional Context for Rare Species Bryophytes and lichens fulfill many important roles in
the ecosystem and are a major component of the floristic bio-diversity of the Olympics. In turn, the OP is of major floristic importance to the region because of its large habitat diversity and because of the occurrence of many species that are known from only a few localities in the PNW (such as Vulpicida tilesii), the conterminous 48 States (such as Bunodophoron melanocarpum, fig. 6), or Washington (such as Splachnum sphaericum, fig. 12). The liverwort Tritomaria exsectiformis (ONHIC list 2; SEIS J2-8a) occurs in a single site (on Olympic National Forest land), which represents the only known site in Washington and Oregon west of the Cascade Range for this species (J. Harpel, oral commun., 2003). The preservation of biological diversity is becoming increasingly important with the advance of anthropogenic habitat conver-sion. It seems inevitable that the main reservoir of diversity is going to be the National Park system and other unmanaged areas. ONP is essentially an unmanaged island in a managed
56 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
landscape. Species associated with old-growth forests in Washington and Oregon, such as the mosses Schistostega pennata (WNHP G4 S2, fig. 13), Tetraphis geniculata (WNHP G5 S2, fig. 14), and Iwatsukiella leucotricha (WNHP G2/3 S1) were once more widespread, and generally only occur in the managed landscape on legacy substrates from the original old-growth forest. Schistostega requires large, shaded, upturned, soil-clad root wads. Tetraphis geniculata requires large, shaded wood debris in an advanced state of decay. After these legacies have been depleted, commercial forests are unlikely to provide a spatially and temporally continuous habi-tat for these species. Their persistence may only be guaranteed in large tracts of old-growth forest, where their microhabitats have greater temporal and spatial continuity.
Although some species of cryptogams seem to disperse easily via light-weight, airborne spores, many spores are not robust to long distance/time travel (Tan and Pocs, 2000). Most species depend on vegetative propagation and slow dispersal through forests (Tan and Pocs, 2000). Consequently, if popu-lations of rare species are lost, they would probably return slowly, if ever. Attempts to revitalize managed landscapes will need to restore the ecological infrastructure and the connectiv-ity to centers of diversity that still remain on the landscape. The large relatively pristine ecosystems in ONP are some of the last few places that still harbor healthy and extensive populations of some of these obligate old-growth associated species. Because of the island-like nature of the OP, species cannot easily recolonize the area once extirpated. A compre-hensive list of ‘special concern species’ for ONP is unavailable as there is insufficient information to adequately assess the fre-quency of many species. Nevertheless, some candidate species can be identified. The macro-lichen Platismatia stenophylla is common in Oregon and other parts of Washington, including the San Juan Islands, and is hence not listed on any State or federal list. However, this distinctive species is known from a few thalli on a single ridge on the northern OP, and hence may represent a critically endangered element of the flora. The southern OP is less well explored, but our limited fieldwork there did not yield additional records of this species. Other examples of species that are apparently rare on the OP, but may be more frequent elsewhere in Washington, are the PNW endemic mosses Bryolawtonia vancouveriensis and Alsia californica and the lichen Ramalina subleptocarpha.
• ThreatsPeople often assume National Parks are in little need
of active management, but human impacts are far reaching and will continue to increase and affect the National Parks. The direct threats are easiest to control and many activities are already regulated or prohibited in National Parks: com-mercial and scientific collection of bryophytes and lichens, construction of human infrastructure such as trails, roads (runoff), parking lots, and buildings. The indirect effects of fire suppression, air pollution, the introduction of invasive exotic organisms, and rapid climate change induced by global
warming all can have major impacts on natural systems in the long term.
Wildfire and fire suppression. In the event of wild-fire, Park managment may attempt to protect critical locations of rare organisms. Periodic fire, on the other hand, maintains habitat diversity and continuity at the landscape level. Low-intensity fires help create open-forest conditions where many lichens and bryophytes thrive. For example, the Blue Mountain fire in ONP eliminated many corticolous lichens, but favored others, including some thriving Bryoria cf. fremontii (fig. 15). A rare PNW endemic lichen, Sulcaria badia (WNHP G? S1), formerly known from a single locality in Washington on the Dungeness Spit, may have gone extinct from the OP as a result of habitat conversion and the effects of fire suppression (Peterson and oth-ers, 1998). Other lichens grow exclusively on charred wood (for instance, Hypocynemyces species), which is still a common sight in many areas of the ONP. However, a policy of fire suppression in National Parks may jeopardize the long-term supply of charred wood.
Air pollution. Air quality is generally good on the OP, but this may change with increasing human population. Pollutants from Asia are now periodically measur-able in ONP (Edmonds and others, 1998), and may increase in the future, eventually to a level where they are harmful to pollution-sensitive lichens. This effect can be monitored in the critically endangered Usnea sphaecelata (WNHP Priority 1, G?S1, fig. 16). It is found in only a few places in Washington, and was recently discovered in ONP. The high fog-bound peaks and ridges favored by this species intercept aerosols, which present a serious threat to this pollution-intol-erant species. Much could be learned from mapping the population of this rare species and seeking other populations on nearby ridges. If the population size is sufficient, pollutant concentrations in its tissue could be monitored periodically along with tissue from com-mon species nearby. Although only an informal survey was conducted, many other rare species were found in the immediate vicinity, making a pollutant monitor-ing effort even more worthwhile. Examples include Alectoria nigricans (Priority 1 G4 S1), A. ochroleuca (Priority 1 G4 S2), Ahtiana sphaerosporella (Priority 2, G? S?), Brodoa oroarctica (Priority 1, G2 S1), Bryoria nitidula (to be verified, possibly new to Washington), Collema undulatum var. granulosum (Washington status unknown), Cornicularia nor-moerica (Priority 2, G? S2), Leptogium minutissimum (Washington status unknown), Leptogium platynum (Washington status unknown), Lobaria linita (Prior-ity 2 G? S?), Nodobryoria subdivergens (Washington status unknown), Umbilicaria cylindrica (Priority 2 G3 S1), U. havaasii (Priority 2 G3 S1), U. kraschenin-nikovii (Priority 2 G3 S1), U. proboscidea (Priority
Discussion 57
Figure 9. Tayloria serrata (composite photograph).
Figure 10. Blepharostoma arachnoideum.
Figure 11. Pseudocyphellaria rainierensis.
Figure 13. Schistostega pennata.
Figure 12. Splachnum sphaericum.
Figure 14. Tetraphis geniculata.
58 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
2 G4S1), and U. rigida (Priority 2 G2 S1, fig. 17). The liverwort Lophozia excisa (Washington status unknown) found at the same site is not known from anywhere else on the OP.
Exotic cryptogams. The introduction of exotic spe-cies of non-vascular plants is not yet a major problem, though it seems inevitable that parks will see them in the future. Orthodontium lineare (fig. 18), a South-ern Hemisphere species, was accidentally imported to Europe, apparently via raw log transports. It has proved an aggressive, invasive species and is now widely distributed in western Europe (Schofield, 1985; van Dort and others, 1998). It may have contributed to the decline of a related species native to England, now critically endangered (http://www.ukbap.org.uk/UKPlans.aspx?ID=485, access restricted, Novem-ber 2005). Another moss, Campylopus introflexus (fig. 19), has spread across western Europe and now is also spreading rapidly in California and Oregon (Christy and Wagner, 1996). In 2001, it was found on the OP for the first time, along the Moclips highway in Grays Harbor County. In 2004 it was found in ONP on Tivoli Island, Lake Ozette. This species grows in thick sods and can replace native vegetation on sandy soils (van Dort and others, 1998). The European species Pseudo-scleropodium purum is one of at least six introduced mosses that are now established in British Columbia (http://srmapps.gov.bc.ca/apps/eswp/, November 2005). It is common in lawns of south coastal British Columbia (Schofield, 1992) and Seattle. It has also been found on the San Juan Islands (Harpel, 1997) and recently in Port Angeles and Quilcene, Washington. It grows in forests of western Europe, but so far has not been detected growing in forested habitats on the OP. Other introduced mosses, liverworts (such as Lunularia cruciata) and lichens (such as Xanthoria parietina and Phaeophyscia rubrapulchra; McCune and Geiser, 1997, and possibly Scoliciosporum sarothamni; Tønsberg, 1995) have established and are spreading in the PNW.
Other exotic species. Exotic species can exert a major influence on pristine ecosystems from another angle as well. White pine blister rust (Cronartium ribicola) has decimated white bark pine (Pinus albicaulis) in ONP and is also taking a toll on western white pine (P. mon-ticola). Species associated with white bark pine also undergo population declines, but these may have gone unnoticed or undocumented. The lichen ‘whitebark candlewax’ (Ahtiana sphaerosporella; WNHP Priority 2, G? S?, fig. 20) was found in the park in only two places, once on dead whitebark pine and once on dead western white pine (Pinus monticola).
Climate change. Humans are influencing ecosystems in major ways. Spatial and temporal patterns of the land-
scape composition and process are very different than those a few hundred years ago due to human activities. Specifically, continuous mosaics of ecosystems have been disrupted so that habitat continuity and con-nectivity have been diminished. This will be of major concern in light of climate change (whether natural or anthropogenic) and the need for species to migrate. Most plants and animals cannot evolve fast enough to adapt apace with the changing conditions, so they must survive by migrating to new habitats. Those species that cannot migrate or adapt along with the rate of environmental change will likely be out-competed by other species moving into their habitats. The cascad-ing effects of rapid climate change could have a major impact on an island-type flora where species may be more easily lost than replaced. Especially if climate is changing at an unprecedented rate, it will be necessary to restore the ecological infrastructure of the landscape to accommodate the migration of species.
Further Work in Olympic National Park
Bryophytes and lichens are an important component of the ecosystem, fulfilling many roles in nutrient and hydrologic cycles and providing nesting material for many animals and birds and housing for numerous invertebrates. Bryophytes and lichens are used as powerful tools in biomonitoring, have commercial value, are a major component of the biodiversity, and in many areas define the aesthetic appearance of ONP. Consequently, there are many kinds of research into ecosystem function that could benefit ONP and the scientific community, but because of the taxonomic focus of our project, this section is devoted to suggestions for additional floristic work.
• Needs for Existing Bryophyte and Lichen Data Complete the identification and verification of difficult
and interesting specimens. It is important that the identity of some of the most interesting taxa be verified, either by taking specimens to a regional herbarium, by requesting reference specimens, or by sending specimens to taxonomic specialists. Specific needs include the following.
ο Chilocyphes gemmiparous (G2 S1), a liverwort col-lected from a high mountain basin. If verified, it would be new to Washington. This species requires work at an herbarium where it can be compared with a specimen, or it could be sent to Dr. D. H. Wagner (Northwest Botanical Institute).
ο There may be two different undescribed Dermato-carpon aff. luridum in ONP. One came from near the Hurricane Ridge Visitor Center; the other specimen came from the park interior and differs from the first. McCune tentatively examined both specimens and recommended that they be sent to Starri Heiðmarsson
Discussion 59
Figure 16. Usnea sphaecelata.
Figure 18. Orthodontium lineare (photographed in The Netherlands).
Figure 20. Ahtiana sphaerosporella.
Figure 19. Campylopus introflexus
(photographed in Coos Bay, Oregon).
Figure 15. Bryoria cf. fremontii.
Figure 17. Umbilicaria rigida.
60 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
(Icelandic Institute of Natural History), a taxonomic expert on Dermatocarpon.
ο There may be an undescribed liverwort in the genus Scapania that was collected from a few localities in ONP. Further work could elucidate whether this species is undescribed, or perhaps is one of the more recently described species from the region. Additional literature will be needed.
Correcting problems in ONP non-vascular plant her-barium. During the past century, independent of this project, some 55 collectors deposited more than 5,000 non-vascular plant collections at ONP. Recently, the outdated nomenclature of many of these collections was revised: the lichen nomencla-ture during this project and the bryophyte nomenclature with support from ONP Cultural Resources Management (CRM) where the collection facility is housed. K. Hutten, with support of CRM, determined 400 specimens with incomplete identi-fications, collected by N. Buckingham. Only specimens with obvious identification errors were re-determined. Other obvi-ous identification errors have been identified, but at present no funds are allocated for work on the historic collections (for example, most if not all D. bonjeanii are probably D. scopar-ium/D. howellii, many Claopodium whippleanum are Pseu-doleskea patens; the determinations in the genus Sphagnum have proved unreliable). Some species concepts have been modified recently, requiring work on all previously determined collections of Pyrrospora cinnabarina for instance, as many pertain to the recently described P. gowardiana (Spribille and Hauck, 2003); similarly, most of the Menegazzia terebrata are M. subsimilis (Bjerke, 2003), several species groups in the lichen genus Fuscopannaria have recently been reworked (Jørgensen, 2000), and so forth.
Generating spatial data from ANCS+ records. Bryophyte and lichen data from other collectors are stored in ANCS+, the NPS-mandated curatorial database. Because of major inadequacies of the spatial component of ANCS+, these his-toric data cannot easily be mapped. With some further effort, however, one could generate coordinates for the roughly 600 different localities where these 5,000+ specimens were collected. These spatial data would allow the addition of thousands of data points to the cryptogam distribution maps of the OP. Geographic and elevational distribution maps also provide a useful tool for identifying outliers that often signal misdeterminations.
Obtain data from other sources. The diverse and abun-dant non-vascular plant flora of ONP has inspired consider-able interest. In the last 30 years, 25 researchers were issued a collecting permit for lichen and bryophyte work in Olympic National Park. Several workers renewed their permits for several years, sometimes accompanied by peer scientists. Apparently, some researchers worked in ONP without a permit or with oral permission, leaving no records at the Park. Although the combined effort was substantial, it left a com-paratively small research legacy at ONP. Many researchers did not deposit voucher specimens at ONP, or deposited few
specimens while making sizeable collections. Many research-ers apparently did not submit an informal collection report, and only one researcher submitted electronic records. Only a limited amount of the gathered data was published in the scientific literature, and few researchers sent copies of their published work to ONP. Therefore, databases are missing data for thousands of specimens collected in ONP. ONP lacks locality data on hundreds of species known from the ONP and OP, including:
ο Dermatocarpon luridum WNHP Priority 1, G? S2 lichen, Clall, Jeff County,
ο Tholurna dissimilis WNHP Priority 1, G? S2 lichen, Hurricane Ridge,
ο Umbilicaria scholanderi WNHP Priority 1, G1 S1 lichen, Mason County,
ο Brachydontium olympicum WNHP G3 S1 moss, Clallam County,
ο Brotherella roellii WNHP G4 S1 PNW endemic moss, Clallam, Jefferson Counties,
ο Sphagnum alaskense newly described moss, not yet listed, Cape Alava Andrus & Janssen 2003,
ο Gymnomitrion ONHIC G5 S2, liverwort, concinnatum park interior Jefferson County? (Christy and Wagner, 1996; fig. 21),
ο and possibly other rare species.Although a portion of these missing specimens may be
adequately cared for in a university or agency collection, the data are effectively inaccessible to ONP, because most facili-ties do not have their herbarium data available electronically. The storage of all inventory data in an appropriate database is a critical component in ONP’s mandate to protect its floris-tic diversity. From such databases, researchers can begin to understand how species are distributed in the park and where rare species are located. Rapid assessments can be made using GIS in the event of environmental or human-induced impacts. ONP could considerably advance its baseline cryptogam data by requesting and compiling data published in the scientific literature, from ONP collections deposited in regional her-baria, from other agencies, and from previous collectors.
• Needed Field Work Geographic areas. The cryptogam diversity encompassed
within the boundaries of ONP is not fully known because there are still many areas that remain unexplored or under-explored by bryologists or lichenologists. Entire river drainages have not been visited (for instance, Bogachiel, Queets, Skokom-ish, Duckabush; fig. 1). The focus of the largely plot-based
Discussion 61
work described here was on specific forest types, therefore many other areas were not surveyed, examples of which are addressed below.
Ecological zones. Some ecological zones have not been adequately surveyed. Significantly, the importance of a high-elevation, hyper-maritime (fog-intercepting) belt to bryophytes and lichens was not fully appreciated until 2002. A USFS project in this zone yielded the first record in Washington of a particularly interesting disjunctive moss, Iwatsukiella leuco-tricha (fig. 22), just outside of the Park. In North America, this arboreal moss is known only from a few locations along the West Coast. It is considered critically endangered in Washing-ton, and was listed WNHP G2/3 S1 in 2004 (J. Harpel, written commun., 2004). In Oregon and Washington, its high-eleva-tion, ocean-influenced habitat has been nearly completely logged. The remaining populations, including some that pre-dictably exist in ONP, are of vital importance for the westward reintroduction of this species, should habitats become suitable again.
Other rare taxa also exist in the hyper-maritime zone. The lichen Bryoria tenuis is new to the OP and possibly Wash-ington (suggested for listing WNHP), and B. bicolor is listed in Oregon. The former Survey and Manage moss Racomitrium aquaticum (WNHP G5 S3, fig. 23), considered rare through-out the PNW, is frequent on fog-intercepting ridges on the OP. It has recently been redescribed as R. ryszardii, and in that interpretation is a PNW endemic. At least four other region-ally rare former Survey and Manage species were frequently encountered in this zone (these occur outside the zone as well), including the endemic lichen Pseudocyphellaria rainie-rensis (WNHP Priority 2, G? S2, fig. 11) and the yet unlisted P. mallota (Tønsberg, 1999b; fig. 7), called “one of our rarest lichens” (http://www.proaxis.com/~mccune/mallota.htm, November 2005). The number and relative abundance of rare taxa in this high-elevation belt suggest that further surveys would be appropriate.
Macrohabitats. Several habitat types remain underex-plored.
ο Mountain ridges provide habitat for many rare species and should be surveyed using an intuitively controlled method (see “Air Pollution” under subsec-tion “Threats” for an example of the number of rare species that may be found on some high-elevation ridges in ONP).
ο Deciduous forests have a diverse lichen and bryophyte flora. Some types that have not been explored during this project include forest types dominated by Acer spp., Alnus sinuata, Arbutus menziesii, Salix spp., Populus trichocarpa, P. tremuloides, or Prunus emarginata.
ο Alnus rubra-dominated riparian and floodplain forests bear diverse lichen floras when old and not too shaded, particularly on nutrient rich soils (Tønsberg, oral com-mun., 2003). Several rare lichens are known from these
habitats, for example Szczawinskia tsugae (WNHP G? S1) and Spilonemella americana (not yet listed, recently described from type locality at the Hoh River).
ο Quercus garryana forests are becoming exceedingly rare on the northern OP. The authors have only exam-ined a polluted fragment, slated for development, near the city center of Sequim. It contains our only collec-tion of the moss Alsia californica, blue listed in BC (S2S3). A recently described lichen species, Physconia fallax, is found on oak trunks and is known from only one other place in Washington (S. Jovan, oral com-mun., 2003). The oak trunks also support Pyrrhospora quernea (WNHP, G? S?; S&M ‘E’). A cedar fencepost in the area supports Thelomma occidentale (WNHP G? S1). The oaks also support a large population of another rare lichen, Ramalina subleptocarpha, on the OP. It reaches the north end of its range here and is listed in British Columbia (S1), but not in Washington. The moss Tortula laevipila var. meridionalis (WNHP G3/G4 T1, S1; J. Harpel, written commun., 2004) was reported by Lawton (1971) from only one locality in North America, on oak in Victoria, British Columbia. It is now known from a few additional localities in the PNW, and it should also be sought in the few oak stands remaining on the Peninsula.
ο The immediate coast needs further surveys. Several rare lichens are exclusively coastal and are expected to occur on the OP coast but have not yet been found (for example, Hypogymnia heterophylla and Heteroder-mia sitchensis). These and several others are expected because they have been found farther north and south along the coast. The critically rare coastal species Heterodermia leucomelos, and Bryoria subcana were recently found on Mt. Walker on USFS land (L. Geiser, written commun., 2003), but their distribution along the OP coast is unknown. Because many of these lichens are arboreal, they are more likely to be found after high winds when freshly fallen branches are com-mon. An opportunistic project after a major wind event would be ideal to assay these lichens. One liverwort that may be found this way on the immediate coast is Scapania scandica (only two records in Washington).
ο Bogs in ONP urgently need detailed surveys to detect and protect rare species. Splachnum ampullaceum (fig. 8) and S. sphaericum (fig. 12) are new to Washington, per this project, and hence are not on the Washington list. The former is known from a single historic locality in Oregon, the latter appears to be new to the PNW. Both species are known from only one coastal bog complex on the OP. Tetraplodon mnioides (WNHP G5 S1; see figure 30) is known from six forested bogs, all outside of the Park, but Christy and Wagner (1996) show a record within ONP (details not known).
62 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
Tayloria serrata (WNHP G4? S1; fig. 9) is a rare PNW species in the Splachnaceae that on the OP is less restricted to bogs than the other species. It has been found in five localities, including an alpine location. Another bog associated moss Tetraplodon cf. angus-tatus (blue listed in BC: G4, S2S3), if verified, would also be new to the PNW. Of the 25 Splachnaceae collections, only two were found during the systematic sampling effort, indicating that these distinctive species are more easily detected by an intuitively controlled sampling method. It is very easy, however, to unwit-tingly damage these populations by trampling and indiscriminate collecting. Any future surveys of ONP coastal bogs should have the important goal of treading especially lightly, and all surveyors should be aware of the fragile and critical habitats of the Splachnaceae.
ο Some mountain and ridge tops are floristically very interesting and may be impacted by view-seeking visi-tors. One of the most frequently visited peaks in ONP supports the only known Washington population of the moss Rhytidium rugosum (G4 S1; fig. 24). Its presence provides support for the glacial refugia hypothesis. Spence (1983) indicated that the population was quite large in 1983, but this population was not relocated during an informal survey in 2000. A detailed survey to relocate and map the population would enable it to be monitored and protected in the future. The lichen Leptogium saturnicum is also restricted to this moun-tain on the OP.
Microhabitats. Bryophytes and lichens often occur on specific substrates or in specific microhabitats. Because such places are not evenly distributed on the landscape, such spe-cies cannot be adequately sampled with a stratified random sampling scheme based on vascular plant associations. Instead, they must be found by seeking specific microhabitats in the field using an intuitively controlled sampling method. Many habitats, microhabitats, and substrates have been under-explored.
ο Areas in and around waterfalls, streams, springs, and other wet areas typically have a diverse bryoflora that differs markedly from nearby floras not influenced by flowing water. Several rare species are known from these habitats. Crumia latifolia (WNHP G3 S1; fig. 25) is a moss that is known from only four places in Washington, three of which are on the OP. Fissidens grandifrons (WNHP G4 S2; fig. 26) is found on rock in seeps or in gentle streams that are always calcium-enriched including several watersheds on the OP. Preissia quadrata (ONHIC List 3, G5 S2) is known from a single locality near a calcium-enriched waterfall in ONP. There are also lichens that are associated with wet areas. Hydrothyria venosa (WNHP G? S2; fig. 27) is known from one alpine wetland in ONP.
ο Rock and rocky outcrops often have stable exposure and moisture regimes on which diverse bryophyte and lichen communities develop. Many rare non-vascular plants are known from such habitats. Pilophorus nigri-caulis (WNHP Priority 1, G?, S2; fig. 28), for example, has recently been found (new to OP list) when survey-ors passed through areas with exposed rock on Colonel Bob and the Skyline trail. Myurella julacea (WNHP Priority 2 G4 S1) is a rare moss that in Washington is only known from rock crevices on the OP. Radula obtusiloba subsp. polyclada is a rare liverwort that has been found on rock walls on the OP. Neither bryophyte has been collected in the past two decades. The authors suspect that these species still occur, but their habitats have not been surveyed.
ο Mineral soil habitats (roadcuts, stream/creek banks, floodplain) are very common but have not been sur-veyed. Discelium nudum (WNHP G5 S1; fig. 29) in Pacific North America occurs on steep (vertical) stream banks in the Hoh and the Bogachiel Rivers and may be under-reported.
ο Animal waste provides a nutrient-enriched growth substrate for a number of non-vascular plants. Several lichens grow on exposed rocks where raptors perch, and a handful of moss species grow exclusively on coyote or elk excrement (and possibly owl pellets; fig. 30). All of these mosses are very rare.
ο Dermatocarpon sp. nov., a lichen possibly new to sci-ence, is known only from natural drainages that now receive runoff from the NPS Visitor Center parking lot. Surveys of similar drainages would ascertain the status of this suspected new lichen species. Additional species of interest occur on soil nearby, including the liverworts Riccia sp., Athalamia hyalina (fig. 31), and Asterella saccata, known only from this location in ONP. The liverworts can only be detected shortly after snowmelt in spring, and may be more widespread than we realize, considering the strong summer bias in the present data. Surveys after snowmelt are recommended to find additional populations of these rare alpine liver-worts in ONP.
ο There are other rare species needing further surveys, but are not mentioned here. For a more exhaustive list, see Appendix A.
Species-specific surveys. There are a number of criti-cal species for which more detailed surveys are particularly needed.
ο Relocate populations of rare species. Several rare species were collected from ONP many years, even decades ago. If general collection localities can be determined, it would be important to attempt to relo-cate populations of these taxa (see examples in section “Obtain data from other sources”).
Discussion 63
Figure 21. Gymnomitrion obtusum is very similar to G. concinnatum and occurs in the same habitat.
Figure 22. Iwatsukiella leucotricha
Figure 24. Rhytidium rugosum (photograph of specimen from Denali National Park, Alaska).
Figure 23. Racomitrium aquaticum.Figure 25. Crumia latifolia.
64 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
ο The only known locality for the lichen Karnefeltia californica is adjacent to an NPS sewage-evaporation field. The species is currently listed in WNHP as G2 S2. The State listing should probably be S1, however, (see “Rare species on the OP”) since there are only five collections in Washington. Many early reports of this species are the more inland species Karnefeltia merrillii. Because the species is also federally listed, a USFS crew has surveyed for this and other rare coastal lichens on the OP as part of a project ranging from California to Washington in 2000. These surveys did not yield further detections of this species (L. Geiser, written commun., 2004). Surveys of adjacent for-est bogs would put the status of this population into perspective in advance of the planned expansion of the sewage facility and the presumed increase in human impacts around the evaporation field.
ο The only known population of the rare moss Rhytidium rugosum may be declining and may be affected by Park visitors (see mountain tops, under macrohabitats). Surveys are needed to locate and assess the population and the potential for impacts.
ο A lichen species in the genus Dermatocarpon, sus-pected to be new to science, was found in a natural drainage area receiving runoff from an NPS parking lot. Further surveys are needed to assess the extent of the population in nearby drainages.
ο The lichen Bunodophoron melanocarpum (fig. 6) was not previously known from the lower 48 States, and it was proposed for listing by WNHP in 2003. A few years after its initial discovery, it is still known from only four or five sites on the OP. Locations near Ozette are the only known locations of this lichen on federal land. The ONP therefore has considerable responsibil-ity for the protection of this species, yet the extent of the population and the existence of other populations is unknown.
ο Splachnum ampullaceum (fig. 8), S. sphaericum (fig. 12), T. mnioides (fig. 30), and T. cf. angustatus are species that occur on elk excrement deposited in coastal bogs. Several species are new to Washington and so far are known only from one coastal bog com-plex on private land. Elk herds may be diminishing in the coastal zone, providing added reason for concern.
ο Sulcaria badia (WNHP G? S1) may be extinct from the Dungeness (Peterson and others, 1998), the only known site in Washington. It is one of the region’s rarest lichens and one of few that has been placed as critically endangered on the global red list (Peterson and others, 1998). Surveys conducted in other suitable areas, in especially the northeast OP, may locate extant populations. In Oregon and California it has been found most often on oak, but also on bigleaf maple,
Douglas fir, Oregon ash, and ponderosa pine (http://www.fs.fed.us/r5/mendocino/projects/hfi/docs/ sulcaria-badia.pdf, November 2005).
ο Brotherella roellii (WNHP G3 S2, Survey and Manage) is a PNW endemic moss that was found in the Dose-wallips decades ago. Relocation efforts have not been successful (J. Harpel, written commun., 2004), but surveys of adjacent ONP may locate more populations.
ο A rare liverwort, Marsupella emarginata var. aquatica (ONHIC G5 S1; S&M ‘B’), was collected in associa-tion with another species from the Park interior. A better specimen and a population survey are needed.
ο Only a few populations of Discelium nudum (WNHP G5 S1; fig. 29) are known from the lower 48 States, two in the Hoh River valley, the other along the Bogachiel River. The specialized sites that this species occupies are often eroded or quickly invaded by vas-cular plants, so the sites are often quite ephemeral. In British Columbia, where the species is red-listed, there are no known currently occupied sites (W. Schofield, written commun., 2004) and the only other site known in Pacific North America is in California (Norris and Shevock, 2004b). Notably, the Pacific Coast popula-tion of Discelium nudum bears sporophytes that are often four to five times the size noted in the rest of its world range. The capsules are often elongate rather than subspheric, and setae can exceed 3 cm, compared to less than 1 cm elsewhere in its range (Schofield, 2003). If the Pacific Coast population is isolated enough to be genetically distinct, the global ranking in the WNHP would increase considerably.
ο The Olympics are suspected to provide good habitat for a number of regionally rare species that have not yet been recorded. Christy and Wagner (1996) and Harpel (oral commun., 2000) predicted that Iwatsukiella leucotricha (fig. 22), one of the region’s rarest species, would likely occur in the Olympics. When the USFS ordered a survey for this species, several populations were detected from areas on the west OP. Although it is not yet known from ONP, it is likely to be there. Other rare species suspected to occur in the Olym-pics include Bartramiopsis lescurii (R. Lesher, oral commun., 2000), Pleuroziopsis ruthenica, and Radula brunnea (Christy and Wagner, 1996; http://www.or.blm.gov/surveyandmanage, temporarily unavailable November 2005).
ο Request specimens and data from future collectors. As mentioned previously (under: Obtain data from Other Sources), valuable information has been lost to ONP or is essentially inaccessible, because many collectors did not deposit voucher specimens at ONP. The authors have been unable to find locations for rare species documented from ONP, and verifying a rare species is
Discussion 65
Figure 30. cf. Tetraplodon mniodes (WNHP G5 S1) on an owl pellet or coyote scat.
Figure 31. Athalamia hyalina.
Figure 29. Discelium nudum.
Figure 27. Hydrothyria venosa inspection by James Walton.
Figure 26. Fissidens grandifrons.
Figure 28. Pilophorus nigricaulis.
66 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
difficult without a voucher specimen. Furthermore, a good reference collection makes specimen identifica-tion easier and increases confidence in identifications. Travel to other herbaria for reference material can be costly, leaving some very interesting specimens unidentified. Moreover, data deposited at the Univer-sity of Washington (WTU) and many other herbaria are not yet entered electronically, making it very time-consuming for ONP staff to locate any quantity of ONP data in the large WTU collection. It may not be necessary to retain every specimen of the most com-mon species at ONP, and certainly not type specimens, but ONP should at least retain those data electronically. North Cascades National Park already requires scien-tists to submit delimited electronic collection data from collectors, relieving curatorial staff from cumbersome data entry. Because data deposited at ONP are entered electronically, they can easily be made available to researchers elsewhere. The ONP collection is particu-larly deficient in crustose lichens, such that they were excluded from the present project because they could not be efficiently identified. Requesting duplicate specimens of crustose lichens from future collectors is an important need for ONP. Considering the impor-tance of having distributional data on any species, collectors should be encouraged to deposit all data at ONP. Storage space at ONP is not a limitation, because a new facility is planned.
ConclusionsThe importance and uniqueness of the non-vascular flora
of ONP are hard to exaggerate. Not only are cryptogams key to the ecosystem function and aesthetic appeal of ONP forests, they represent a biogeographically significant array of rare and endemic species and a sizable component of the Park’s biodiversity. This project has made important steps toward a comprehensive survey of these organisms. It has put the Park’s resources in a regional context by documenting rare species, some for the first time in the conterminous United States, and a few species that are apparently undescribed. ONP has healthy populations of some species that are rare outside of the Park, where habitats have been destroyed. This data set can form the basis for further research on community stucture and successional patterns, and could be used as a tool for eventual restoration efforts outside of the Park.
Forests are one of the most important habitats for bryo-phytes and lichens globally, and conservation of forests is key to maintaining their biodiversity (Tan and Pocs, 2000). The NPS has had a long-standing mandate to protect habitats and a more recent mandate to inventory its resources so it can better protect biodiversity (National Park Service, 2001). Conserva-tion of bryophytes and lichens worldwide has usually been a
by-product of saving habitat for larger flagship, charismatic, or keystone species (Tan and Pocs, 2000). ONP is one of few places in the world where bryophytes and lichens, collectively, are conspicuous flagship organisms themselves. Consequently, ONP has the opportunity to bring them to the attention of the public, government, and other conservation agencies. The inventory described here is a tool to help ONP understand and conserve its non-vascular resources.
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70 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
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ELLO
W G
5 S3
S4
Myu
rella
jula
cea
G4
S1Y
ELLO
W G
5 S3
S5
Onc
opho
rus w
ahle
nber
gii
G5
S1Y
ELLO
W G
5 S3
S4
Orth
othe
cium
chr
yseu
m
G4
S1Y
ELLO
W G
5? S
3S5
Orth
otric
hum
affi
ne
BLU
E G
3G5
S2S3
Orth
otric
hum
alp
estre
BL
UE
G4G
5 S2
S3
Orth
otric
hum
pel
luci
dum
Li
st 3
G4
S1Y
ELLO
W G
3G5
S3S 4
Orth
otric
hum
pyl
aisi
i G
5 S1
BLU
E G
4G5
S1S3
Orth
otric
hum
rivu
lare
R
ED G
4 S1
Philo
notis
font
ana
var.
amer
ican
aBL
UE
G5T
NR
S2S3
Philo
notis
font
ana
var.
pum
ilaBL
UE
G5T
4T5
S2S3
Philo
notis
yez
oana
R
ED G
2G3
S2
Phys
com
itriu
m p
yrifo
rme
BLU
E G
5 S2
S3
Plag
iom
nium
cili
are
BLU
E G
5 S1
S3
Plag
iopu
s oed
eria
na
G5
S2Y
ELLO
W G
5? S
3S4
Plat
yhyp
nidi
um ri
pario
ides
G
4 S1
List
3 G
4 S1
91
72 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Pohl
ia a
tropu
rpur
ea
BLU
E G
4G5
S2S3
Pohl
ia b
olan
deri
BLU
E G
U S
2S3
Pohl
ia c
ampt
otra
chel
aBL
UE
G3G
5 S1
S3
Pohl
ia c
olum
bica
G
2 S1
BLU
E G
3G5
S1S3
Pohl
ia e
long
ata
BLU
E G
4G5
S2S3
Pohl
ia e
rect
a G
3 S1
RED
G3G
5 S1
Pohl
ia fi
lum
BL
UE
G4G
5 S2
S3
Pohl
ia le
scur
iana
BL
UE
G4?
S1S
3
Pohl
ia lu
dwig
ii BL
UE
G5?
S2S
3
Pohl
ia o
btus
ifolia
BL
UE
G2G
4 S2
S3
Pohl
ia p
acifi
ca
BLU
E G
U S
1S3
Pohl
ia v
exan
s BL
UE
G3G
5 S2
S3
Poly
trich
um lo
ngis
etum
BL
UE
G5
S2S3
Poly
trich
um se
xang
ular
e Li
st 3
G4
S1Y
ELLO
W G
4 S3
S5
Poly
trich
um st
rictu
m
G5
S2Li
st 2
G4
S1Y
ELLO
W G
5 S3
S5
Rac
omitr
ium
aqu
atic
um
G5
S3E
EG
3Q N
3Q S
2Li
st 3
Y
ELLO
W G
2G4
S3S 5
1,3
ER
acom
itriu
m m
icro
carp
on
G5
S1R
acom
itriu
m p
acifi
cum
Li
st 2
BL
UE
G3
S3
Rac
omitr
ium
pyg
mae
um
BLU
E G
U S
1S3
Rha
bdow
eisi
a cr
ispa
ta
BLU
E G
5 S2
S3
Rhi
zom
nium
nud
um
X-W
A, B
-OR
(CA
)B-
OR
(CA
)G
4 N
4 S4
List
3 G
4 S2
YEL
LOW
G3G
5 S3
S 51,
3B
As
SS-O
Rhi
zom
nium
pun
ctat
um
BLU
E G
5 S1
S3
Rhy
tidiu
m ru
gosu
m G
4 S1
List
2 G
5 S1
YEL
LOW
G5
S3S4
Schi
stid
ium
aga
ssiz
ii BL
UE
G3G
5 S2
S3
Schi
stos
tega
pen
nata
G
4 S2
AA
G3G
4 N
3N4
S3Li
st 2
G4
S2Y
ELLO
W G
4 S3
S41,
3A
As
SS
Scle
ropo
dium
tour
etii
var.
colp
ophy
llum
BLU
E G
4TN
R S1
S3
Spha
gnum
qui
nque
fariu
m
BLU
E G
5 S2
S3
Spla
chnu
m a
mpu
llace
um
List
2 G
4 S1
YEL
LOW
G4
S3S5
Spla
chnu
m v
ascu
losu
m
BLU
E G
3G5
S1S3
Tayl
oria
serra
ta
G4
S1Li
st 2
G4
S1Te
traph
is g
enic
ulat
a G
4 S2
AA
G3
N3
S3Li
st 2
G4
S1Y
ELLO
W G
3G5
S3S 5
1,3
PBA
As
SS
Tetra
plod
on a
ngus
tatu
s BL
UE
G4
S2S3
Tetra
plod
on m
nioi
des
G5
S1Li
st 2
G4
S1Y
ELLO
W G
4 S3
S4
Tham
nobr
yum
nec
kero
ides
G
3 S2
List
3 G
4 S2
YEL
LOW
GU
S3S
5
Tim
mia
meg
apol
itana
BLU
E G
5 S2
S3
92
Appendix A 73
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Tortu
la a
mpl
exa
BLU
E G
2G4
S2S3
Tortu
la b
olan
deri
RED
G3G
5 S1
Tortu
la su
bula
ta
List
3 G
5 S1
BLU
E G
5? S
2S3
Trip
tero
clad
ium
leuc
ocla
dulu
m
List
2 G
3 S2
BLU
E G
3 S2
S3
(Tot
al 1
00 M
OSS
ES)
LIV
ERW
OR
TSA
nast
roph
yllu
m m
inut
umLi
st 3
G4
S1B
arbi
loph
ozia
lyco
podi
oide
s Li
st 2
G5
S1C
alyp
ogei
a sp
hagn
icol
a Li
st 2
G4
S2D
iplo
phyl
lum
alb
ican
s X
X e
ffect
ive
G5
N4
S41,
3D
iplo
phyl
lum
plic
atum
B
BG
4 N
3 S2
List
2 G
4 S2
1,2
BA
s
Gym
nom
itrio
n co
ncin
natu
m
List
2 G
5 S2
Har
pant
hus f
loto
vian
us
List
3 G
5 S1
Her
bertu
s adu
ncus
E
EG
5 N
3N4
S1Li
st 2
G5
S21,
3A
s
Jam
eson
iella
aut
umna
lis
List
3
Loph
ozia
gill
man
ii Li
st 3
G5
S1M
arsu
pella
con
dens
ata
List
3 G
3 S1
Mar
supe
lla e
mar
gina
ta v
ar. a
quat
ica
BB
G5T
3 N
3 Li
st 2
G5T
3 S1
1,2
BA
s
Mar
supe
lla sp
arsi
folia
Li
st 3
G3
S1M
etzg
eria
tem
pera
ta
List
2 G
4 S1
Nar
dia
japo
nica
Li
st 3
G5
S2Pr
eiss
ia q
uadr
ata
List
3 G
5 S2
Ptili
dium
pul
cher
rimum
Li
st 2
G5
S1R
adul
a ob
tusi
loba
subs
p. p
olyc
lada
Scap
ania
obs
cura
Li
st 3
G3
S1Sc
hofie
ldia
mon
ticol
a Li
st 3
G3
S1Tr
itom
aria
exs
ectif
orm
is
BB
G5
N4
S2Li
st 2
G5
S21,
2B
As
Trito
mar
ia q
uinq
uede
ntat
a B
BG
5 N
4 S1
List
2 G
5 S1
1,3
BA
s
(Tot
al 2
2 LI
VERW
ORT
S)
LIC
HEN
SA
htia
na p
allid
ula
Prio
rity
2 G
? S?
Aht
iana
spha
eros
pore
lla
Prio
rity
2 G
? S?
Ale
ctor
ia la
ta
Prio
rity
1 G
? S2
93
74 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Ale
ctor
ia n
igric
ans
Prio
rity
1 G
4 S2
Ale
ctor
ia o
chro
leuc
a Pr
iorit
y 1
G4
S1A
lect
oria
van
couv
eren
sis
S1
Arth
onia
stel
laris
Pr
iorit
y 2
G?
S?A
rthro
rhap
his c
itrin
ella
Pr
iorit
y 2
G?
S?B
rodo
a or
oarc
tica
Prio
rity
1 G
3 S1
Bry
oria
bic
olor
Li
st 3
G4
S1B
ryor
ia fr
iabi
lis
Prio
rity
1 G
3 S1
Bry
oria
pik
ei
Prio
rity
1 G
2 S2
Bry
oria
pse
udoc
apill
aris
A
AG
1G2
N1N
2 S1
List
2 G
2? S
11,
3B
SS?
Bry
oria
subc
ana
BB
G2G
4 N
2N3
S1Li
st 3
G4
S11,
3B
Bry
oria
tortu
osa
Prio
rity
2 G
2 S3
A-W
A O
P +
1A
-WA
OP
+1
G5
N5
S31,
3A
-WA
OP
+1
Bue
llia
oida
lea
Prio
rity
2 G
? S2
EE
G3?
N3?
S1
List
3 G
4 S1
1,3
EB
unod
opho
ron
mel
anoc
arpu
m
e d a
s new
WN
HP
Cal
iciu
m a
biet
inum
Pr
iorit
y 2
G?
S?B
BG
4G5
N4
S2S3
List
3
4B
Cal
iciu
m g
lauc
ellu
m
Prio
rity
2 G
? S?
FF
G4G
5 N
4 S4
4F
Cet
relia
cet
rario
ides
Pr
iorit
y 2
G?
S3E
EG
4G5
N4
S2Li
st 3
4
ESS
-WA
?
Cha
enot
heca
ferr
ugin
ea
Prio
rity
2 G
? S?
BB
G4G
5 N
4 S4
List
4
4B
Cha
enot
heca
furf
urac
ea
Prio
rity
2 G
? S?
FF
G4G
5 N
4 S3
List
3
4F
Cla
dina
por
tent
osa
subs
p. p
acifi
caPr
iorit
y 2
G?
S2C
lado
nia
norv
egic
a Pr
iorit
y 2
G3
S2B
BG
4G5
N2N
3 S2
List
3
3B
Coc
cotre
ma
poci
llariu
m
Prio
rity
1 G
2 S1
Col
lem
a fe
cund
um
S2
Col
lem
a fu
rfur
aceu
m
S2
Cor
nicu
laria
nor
moe
rica
Prio
rity
1 G
? S2
Cys
toco
leus
ebe
neus
Pr
iorit
y 2
G?
S1D
endr
isco
caul
on sp
.Pr
iorit
y 1
G?
S1A
-WA
+3
A-W
A +
3 N
4Q S
2Li
st 2
G3
S11,
3B
SS-W
A?
Der
mat
ocar
pon
lurid
um
Prio
rity
1 G
? S2
EE
G4G
5 N
4 S1
S2Li
st 3
1,
3B
SS?
Der
mat
ocar
pon
mou
linsi
i Pr
iorit
y 1
G?
S?Er
iode
rma
sore
diat
um
Prio
rity
1 G
? S1
-Li
st 2
G3
S1 S
11,
3X
SS?
Fusc
opan
naria
lace
ratu
la S
2
Fusc
opan
naria
pac
ifica
G5?
N5
S4S4
Fusc
opan
naria
saub
inet
ii F
FG
3G5
N2
S1?
4F
Gya
lideo
psis
ana
stom
osan
s Pr
iorit
y 2
G?
S1H
alec
ania
viri
desc
ens
Prio
rity
1 G
? S1
94
Appendix A 75
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Het
erod
erm
ia le
ucom
ela
- S
21,
3X
SS?
Hyd
roth
yria
ven
osa
Prio
rity
1 G
? S2
- S
21,
3X
Hyp
ogym
nia
dupl
icat
a Pr
iorit
y 2
G?
S3C
CG
4 N
4 S3
List
3
1,2,
3A
SS-O
R?
Hyp
ogym
nia
ocea
nica
Pr
iorit
y 2
G?
S1X
-Li
st 2
1,
3F
Hyp
ogym
nia
vitta
ta
Prio
rity
2 G
? S?
EE
G4G
5 N
3 SN
A S
?3
EK
aern
efel
tia c
alifo
rnic
a Pr
iorit
y 2
G2
S2-
S2
1,3
XLe
iode
rma
sore
diat
um
-Li
st 2
G3
S1 S
11,
3X
SS?
Lept
ogiu
m b
rebi
sson
ii -
S1
1,3
XSS
?
Lept
ogiu
m c
yane
scen
s A
AG
5 N
4 S1
List
3
S1
4A
SS?
Lept
ogiu
m p
laty
num
S
2
Lept
ogiu
m p
olyc
arpu
m
S1
Lept
ogiu
m te
retiu
scul
um
Prio
rity
2 G
? S?
EE
G4G
5 N
4 Li
st 3
4
ELo
baria
hal
lii
Prio
rity
2 G
? S?
-1,
3X
Loba
ria li
nita
Pr
iorit
y 2
G?
S?A
WC
S S
noqu
alm
ie P
G4G
5 N
3 S3
List
2 G
4 S1
1,2,
3A
SS-O
R?
Loba
ria si
lvae
-vet
eris
Prio
rity
1 G
? S1
S1
Loxo
spor
a el
atin
a Pr
iorit
y 2
G?
S?Lo
xosp
orop
sis c
oral
lifer
a Pr
iorit
y 2
G?
S?-
1,3
XM
icro
lych
nus e
pico
rtici
s Pr
iorit
y 2
G?
S1N
ephr
oma
bellu
m
E-W
A (O
P) +
4E-
WA
(OP)
+4
G3G
5 N
3N4
S24
FSS
-WA
?
Nep
hrom
a oc
cultu
m
Prio
rity
1 G
? S1
AA
G3
N3
S1Li
st 3
G3
S31,
3B
SS?
Nie
bla
ceph
alot
a Pr
iorit
y 1
G?
S1A
AG
1G3
N2
S1Li
st 3
G4
S2 S
11,
3A
SS?
Ope
grap
ha so
redi
ifera
Pr
iorit
y 2
G?
S1O
phio
parm
a la
ppon
ica
Prio
rity
1 G
? S1
Pann
aria
rubi
gino
sa
EE
G4G
5 N
4 S1
List
2 G
4 S1
S2
1,3
ESS
?
Parm
otre
ma
chin
ense
S
1
Parm
otre
ma
crin
itum
S
2
Pelti
gera
lepi
doph
ora
Prio
rity
1 G
? S1
Pelti
gera
pac
ifica
E
EG
3 N
2N3
S2?
4E
SS?
Pilo
phor
us n
igric
aulis
Pr
iorit
y 1
G?
S2-
List
2 G
4 S2
S1
1,3
XSS
?
Plat
ism
atia
lacu
nosa
Pr
iorit
y 2
G?
S3C
(exc
ept O
R C
oast
RG
3G4
N3
S2Li
st 3
4
CSS
-WA
?
Pseu
docy
phel
laria
cro
cata
Pr
iorit
y 2
G?
S3-
4X
Pseu
docy
phel
laria
mal
lota
Li
st 2
G4
S1Ps
eudo
cyph
ella
ria p
erpe
tua
BB
G2G
4 N
2 1,
3B
Pseu
docy
phel
laria
rain
iere
nsis
Pr
iorit
y 2
G?
S2A
AG
3G4
N3N
4 S3
List
3 G
3 S1
S1
1,2,
3A
SS?
Pyrrh
ospo
ra q
uern
ea
Prio
rity
2 G
? S?
EE
G4
N4
S1S2
List
3
1,3
ESS
?
95
76 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Pyrrh
ospo
ra su
bcin
naba
rina
Prio
rity
2 G
? S1
Ram
alin
a m
enzi
esii
Prio
rity
2 G
? S3
Ram
alin
a po
llina
ria
Prio
rity
2 G
? S?
EE
G4G
5 N
4 S1
List
2
3E
SS?
Ram
alin
a su
blep
toca
rpha
S
1
Ram
alin
a th
raus
ta
Prio
rity
1 G
? S1
AA
G3G
4 N
3 S1
4A
Solo
rina
sacc
ata
Prio
rity
2 G
? S?
Stic
ta b
eauv
oisi
i s.l.
Prio
rity
2 G
? S?
-4
XSu
lcar
ia b
adia
Pr
iorit
y 1
G?
S1Li
st 2
G2
S2Sz
czaw
insk
ia ts
ugae
Pr
iorit
y 2
G?
S1Th
elom
ma
occi
dent
ale
Prio
rity
2 G
? S1
Thol
urna
dis
sim
ilis
Prio
rity
1 G
? S2
BB
sout
h of
Col
R.
G3G
5 N
3 S2
List
3 G
4 S1
1,3
BSS
?
Um
bilic
aria
ang
ulat
a Pr
iorit
y 2
G2
S2U
mbi
licar
ia a
rctic
a Pr
iorit
y 2
G3
S1U
mbi
licar
ia c
ylin
dric
a Pr
iorit
y 2
G3
S1U
mbi
licar
ia d
ecus
sata
Pr
iorit
y 2
G3
S1U
mbi
licar
ia h
avaa
sii
Prio
rity
2 G
3 S1
Um
bilic
aria
kra
sche
ninn
ikov
ii Pr
iorit
y 2
G3
S1U
mbi
licar
ia la
mbi
i Pr
iorit
y 1
G?
S1U
mbi
licar
ia ly
ngei
Pr
iorit
y 2
G3
S1U
mbi
licar
ia p
olyr
rhiz
aPr
iorit
y 2
G1
S1U
mbi
licar
ia p
robo
scid
ea
Prio
rity
2 G
4 S1
Um
bilic
aria
rigi
da
Prio
rity
2 G
2 S1
Um
bilic
aria
scho
land
eri
Prio
rity
2 G
1 S1
Um
bilic
aria
vel
lea
Prio
rity
2 G
3 S2
Usn
ea h
espe
rina
EE
G4G
5 N
4 S1
?Li
st 3
G4
S31,
3B
Usn
ea lo
ngis
sim
aPr
iorit
y 2
G?
S2F-
WA
+2
F-W
A +
2G
3G4
N3
S2Li
st 3
4
F-W
A +
2SS
?
Usn
ea ru
bicu
nda
List
3 G
4 S2
Usn
ea sp
hace
lata
Pr
iorit
y 1
G?
S1Li
st 3
G4
S1U
snea
wirt
hii
Prio
rity
2 G
2 S2
Vul
pici
da ti
lesi
i Pr
iorit
y 1
G4
S1
(Tot
al 1
03 L
ICHE
NS)
96
Appendix A 77
App
endi
x A
—co
ntin
ued
(Foo
tnot
es o
n pa
ge 7
8.)
Sp
ecia
l Sta
tus
W
NHP
SM
200
2 AS
RSM
200
3 AS
R
SM
ratin
g
ONH
P
B
C ra
ting
Scie
ntifi
c N
ame
RO
D S
M 2
001
SEIS
20
05
BLM
2005
U
SFS
R6:
O
R/W
A1
23
45
67
8
910
Inde
xes c
reat
ed o
ver a
ll re
cent
199
8- c
urre
nt d
ata
(USG
S/O
NP
data
M. H
utte
n pe
rson
al d
ata)
, but
exc
ludi
ng A
NCS
dat
a be
caus
e of
seve
ral p
robl
ems a
ssoc
iate
d w
ith
that
dat
aset
. Th
ese
data
are
pro
vide
d as
an
indi
cato
r of t
he re
lativ
e fr
eque
ncy
of th
e sp
ecie
s col
lect
ed.
The
OP
loca
litie
s cou
nt is
gen
erat
ed fr
om u
niqu
e sit
es o
nly,
i.e.
du
plic
ate
colle
ctio
ns fr
om a
sing
le si
te a
re e
xclu
ded.
Suc
h un
ique
site
s how
ever
may
occ
asio
nally
be
in c
lose
pro
xim
ity to
eac
h ot
her.
Not
e th
at th
e W
ashi
ngto
n N
atur
al H
erita
ge P
rogr
am h
as n
ot y
et in
clud
ed li
verw
orts
in th
eir l
istin
gs.
Abb
revi
atio
ns u
sed:
W
NH
P= W
ashi
ngto
n N
atur
al H
erita
ge P
rogr
amG
= in
dica
tor o
f glo
bal s
tatu
sS=
indi
cato
r of s
tate
stat
us1=
crit
ical
ly im
peril
ed b
ecau
se o
f ext
rem
e ra
rity
or b
ecau
se it
is p
artic
ular
ly v
ulne
rabl
e to
ext
inct
ion
or e
xtirp
atio
n; ty
pica
lly 5
or f
ewer
occ
urre
nces
2= Im
peril
ed b
ecau
se o
f rar
ity o
r bec
ause
it is
vul
nera
ble
to e
xtin
ctio
n or
ex
tirpa
tion;
typi
cally
6 to
20
occu
rren
ces
3= E
ither
ver
y ra
re a
nd lo
cal t
hrou
ghou
t its
rang
e or
foun
d lo
cally
(eve
n ab
unda
ntly
) in
a re
stric
ted
rang
e;
typi
cally
21
to 1
00 o
ccur
renc
es.
4= A
ppar
ently
secu
re; t
ypic
ally
mor
e th
an 1
00 o
ccur
renc
es5=
Dem
onstr
ably
wid
espr
ead,
abu
ndan
t and
secu
re.
U=
Stat
us u
ncer
tain
; add
ition
al in
form
atio
n is
need
ed.
?= A
dditi
onal
info
rmat
ion
need
ed
ON
HP
= O
rego
n N
atur
al H
erita
ge P
rogr
am:
1 =
Crit
ical
ly im
peril
ed b
ecau
se o
f ext
rem
e ra
rity
(5 o
r few
er o
ccur
renc
es, o
r ve
ry fe
w re
mai
ning
indi
vidu
als)
, or b
ecau
se o
f som
e fa
ctor
of i
ts bi
olog
y m
akin
g it
espe
cial
ly v
ulne
rabl
e to
ext
inct
ion.
2
= Im
peril
ed b
ecau
se o
f rar
ity (6
-20
occu
rren
ces)
, or b
ecau
se o
f oth
er fa
ctor
sde
mon
strab
ly m
akin
g it
very
vul
nera
ble
to e
xtin
ctio
n th
roug
hout
its r
ange
.3
= E
ither
ver
y ra
re a
nd lo
cal t
hrou
ghou
t its
rang
e, o
r fou
nd lo
cally
(eve
nab
unda
ntly
at s
ome
of it
s loc
atio
ns) i
n a
restr
icte
d ra
nge,
or v
ulne
rabl
e to
extin
ctio
n th
roug
hout
its r
ange
bec
ause
of o
ther
fact
ors;
in th
e ra
nge
of
21-1
00 o
ccur
renc
es.
4 =
App
aren
tly se
cure
, tho
ugh
it m
ay b
e qu
ite ra
re in
par
ts of
its r
ange
, es
peci
ally
at t
he p
erip
hery
.
SM=
Surv
ey a
nd M
anag
e th
e pr
ogra
m im
plem
entin
g th
e RO
D=R
ecor
d of
Dec
ision
, app
licab
le to
For
est S
ervi
ce a
nd B
urea
u of
Lan
d M
anag
emen
t for
est m
anag
emen
t in
rang
e of
the
Nor
ther
n Sp
otte
d O
wl.
In
subs
eqen
t A
men
dmen
ts m
ade
to th
e RO
D, r
evisi
ons w
ere
mad
e to
the
statu
s of R
OD
spec
ies
(USD
A a
nd U
SDI 2
001)
. Th
e 20
02 a
nd 2
003
Ann
ual S
peci
es R
evie
w (A
SR) r
esul
ts ar
e lis
ted
in th
e ta
ble
abov
e.
97
78 Inventory of the Mosses, Liverworts, Hornworts, and Lichens of Olympic National Park, Washington: Species List
G =
Indi
cato
r of g
loba
l sta
tus
S
= In
dica
tor o
f sta
te st
atus
1 =
Crit
ical
ly im
peril
ed b
ecau
se o
f ext
rem
e ra
rity
or b
ecau
se it
is p
artic
ular
ly v
ulne
rabl
e to
extin
ctio
n or
ext
irpat
ion;
typi
cally
5 o
r few
er o
ccur
renc
es
2
= Im
peril
ed b
ecau
se o
f rar
ity o
r bec
ause
it is
vul
nera
ble
to e
xtin
ctio
n or
ext
irpat
ion;
ty
pica
lly 6
to 2
0 oc
curr
ence
s
3 =
Eith
er v
ery
rare
and
loca
l thr
ough
out i
ts ra
nge
or fo
und
loca
lly (e
ven
abun
dant
ly) i
n a
rest
ricte
d ra
nge;
typi
cally
21
to 1
00 o
ccur
renc
es.
4
= A
ppar
ently
secu
re; t
ypic
ally
mor
e th
an 1
00 o
ccur
renc
es
5
= D
emon
stra
bly
wid
espr
ead,
abu
ndan
t and
secu
re.
U
= S
tatu
s unc
erta
in; a
dditi
onal
info
rmat
ion
is n
eede
d.
?
= A
dditi
onal
info
rmat
ion
need
ed
N
ote
that
the
WN
HP
has n
ot y
et in
clud
ed li
verw
orts
in th
eir l
istin
gs.
2 ON
HP
= O
rego
n N
atur
al H
erita
ge P
rogr
am
1
= C
ritic
ally
impe
riled
bec
ause
of e
xtre
me
rarit
y (5
or f
ewer
occ
urre
nces
, or v
ery
few
re
mai
ning
indi
vidu
als)
, or b
ecau
se o
f som
e fa
ctor
of i
ts b
iolo
gy m
akin
g it
espe
cial
ly
vuln
erab
le to
ext
inct
ion.
2 =
Impe
riled
bec
ause
of r
arity
(6–2
0 oc
curr
ence
s), o
r bec
ause
of o
ther
fact
ors d
emon
stra
bly
mak
ing
it ve
ry v
ulne
rabl
e to
ext
inct
ion
thro
ugho
ut it
s ran
ge.
3
= Ei
ther
ver
y ra
re a
nd lo
cal t
hrou
ghou
t its
rang
e, o
r fou
nd lo
cally
(eve
n ab
unda
ntly
at s
ome
of it
s loc
atio
ns) i
n a
rest
ricte
d ra
nge,
or v
ulne
rabl
e to
ext
inct
ion
thro
ugho
ut it
s ran
ge
beca
use
of o
ther
fact
ors;
in th
e ra
nge
of 2
1–10
0 oc
curr
ence
s.
4
= A
ppar
ently
secu
re, t
houg
h it
may
be
quite
rare
in p
arts
of i
ts ra
nge,
esp
ecia
lly a
t the
pe
riphe
ry.
3 RO
D =
Rec
ord
of D
ecis
ion;
app
licab
le to
For
est S
ervi
ce a
nd B
urea
u of
Lan
d M
anag
emen
t for
est
man
agem
ent i
n ra
nge
of th
e N
orth
ern
Spot
ted
Ow
l.
1
= M
anag
e K
now
n Si
tes
2
= Su
rvey
Prio
r to
Gro
und-
Dis
turb
ing
Spec
ies
3
= Ex
tens
ive
Surv
eys
4
= G
ener
al R
egio
nal S
urve
ys
4 SM
= S
urve
y an
d M
anag
e; t
he p
rogr
am im
plem
entin
g th
e R
OD
was
am
ende
d se
vera
l tim
es, a
nd
revi
sion
s wer
e m
ade
to th
e st
atus
of R
OD
spec
ies
(USD
A a
nd U
SDI 2
001)
. In
2001
a S
uppl
emen
tal
Envi
ronm
enta
l Im
pact
Sta
tem
ent (
SEIS
) was
rele
ased
.
A =
Rar
e, P
re-D
istu
rban
ce S
urve
ys P
ract
ical
B =
Rar
e, P
re-D
istu
rban
ce S
urve
ys N
ot P
ract
ical
C =
Unc
omm
on, P
re-D
istu
rban
ce S
urve
ys P
ract
ical
D =
Unc
omm
on, P
re-D
istu
rban
ce S
urve
ys N
ot P
ract
ical
or N
ot N
eces
sary
E
= R
are,
Sta
tus U
ndet
erm
ined
F =
Unc
omm
on o
r Con
cern
for P
ersi
sten
ce U
nkno
wn,
Sta
tus U
ndet
erm
ined
X =
Spe
cies
rem
oved
from
list
5 2
002
Surv
ey A
nd M
anag
e Ann
ual S
peci
es R
evie
w (A
SR) r
atin
g (s
ee 4
)
+1:
A –
WA
OP,
WA
W L
owla
nds,
WA
W C
asca
des,
OR
W C
asca
des,
OR
Coa
st R
ange
, OR
Will
amet
te V
alle
y, a
nd C
A C
oast
Ran
ge
D –
WA
& O
R
E –
Cas
cade
s, O
R &
CA
Kla
mat
h, &
CA
Cas
cade
s
+2:
F –
WA
, OR
,
A
– C
A,O
R (C
urry
, Jos
ephi
ne, J
acks
on)
+3
: A
– W
A,
E –
CA
,
E
– O
R (C
oos,
Dou
glas
, Cur
ry, J
osep
hine
, Jac
kson
)
A
– O
R re
mai
nder
+4:
E –
WA
(W-C
asca
des o
utsi
de G
PNF)
,
E
– C
asca
des,
OP)
E
– O
R (K
lam
ath,
Will
amet
te V
alle
y &
Eas
t Cas
cade
s)
X –
Rem
oved
els
ewhe
re
6 200
3 Su
rvey
And
Man
age A
nnua
l Spe
cies
Rev
iew
dat
a (s
ee 5
) 7 2
005
Bur
eau
of L
and
Man
agem
ent (
BLM
) rat
ings
SS =
Sen
sitiv
e Sp
ecie
s
BS
= B
urea
u Se
nsiti
ve
8 Uni
ted
Stat
es F
ores
t Ser
vice
(USF
S) S
peci
al S
tatu
s Spe
cies
Rat
ing
SS =
Sen
sitiv
e Sp
ecie
s 9 S
urve
y an
d M
anag
e R
atin
g (s
ee W
NH
P ra
tings
for e
xpla
natio
n of
the
ratin
g sy
stem
) 10
Brit
ish
Col
umbi
a (G
owar
d 19
96):
1 =
Crit
ical
ly im
peril
ed b
ecau
se o
f ext
rem
e ra
rity
(5 o
r few
er o
ccur
renc
es, o
r ver
y fe
w
rem
aini
ng in
divi
dual
s), o
r bec
ause
of s
ome
fact
or o
f its
bio
logy
mak
ing
it es
peci
ally
vuln
erab
le to
ext
irpat
ion
or e
xtin
ctio
n.
2
= Im
peril
ed b
ecau
se o
f rar
ity (6
to 2
0 ex
tant
occ
urre
nces
or v
ery
few
rem
aini
ng in
divi
dual
s),
or b
ecau
se o
f oth
er fa
ctor
s dem
onst
rabl
y m
akin
g it
vuln
erab
le to
ext
irpat
ion
or
ext
inct
ion.
1 WN
HP=
Was
hing
ton
Nat
ural
Her
itage
Pro
gram
Hutten, M., W
oodward, A. &
Hutten, K.—Inventory of the M
osses, Liverworts, Hornw
orts, and Lichens of Olympic N
ational Park, Washington: Species List—
Scientific Investigations Report 2005-5240
Printed on recycled paper