J Hattori Bot. Lab. No. 92: 91- 123 (Aug. 2002)

BRYOPHYTES FROM TUXEDNI WILDERNESS AREA, ALASKA

I 2 3 W B. SCHOFIELD , S. S. TALBOT AND S. L. TALBOT

ABSTRACT. The bryoflora of two small maritime islands, Chisik and Duck Island (2,302 ha), com­ prising Tuxedni Wilderness in western lower Cook Inlet, Alaska, was examined to determine species composition in an area where no previous collections had been reported. The field study was con­ ducted from sites selected to represent the totality of environmental variation within Tuxedni Wilder­ ness. Data were analyzed using published reports to compare the bryophyte distribution patterns at three levels, the Northern Hemisphere, North America, and Alaska. A total of 286 bryophytes were identified: 230 mosses and 56 liverworts. Bryum miniatum, Dichodontium olympicum, and Or­ thotrichum pal/ens are new to Alaska. The annotated list of species for Tuxedni Wilderness expands the known range for many species and fills distribution gaps within Hulten's Central Pacific Coast district. Compared with bryophyte distribution in the Northern Hemisphere, the bryoflora of Tuxedni Wilderness primarily includes taxa ofboreal (61%), montane (13%), temperate (11%), arctic-alpine (7%), cosmopolitan (7%), distribution; 4% of the total moss flora are North America endemics. A brief summary of the botanical exploration of the general area is provided, as is a description of the bryophytes present in the vegetation and habitat types of Chisik and Duck Islands.

KEy WORDS: Alaska, Tuxedni Wilderness, boreal zone, moss, liverwort, phytogeography.

INTRODUCTION Tuxedni Wilderness Area on the western shore of lower Cook Inlet, Alaska, includes Chisik and Duck Islands (Fig. 1). As part of the Alaska Maritime National Wildlife Refuge managed by the U.S. Fish and Wildlife Service, Tuxedni Wilderness is a Class I air quality area (Clean Air Act, 42 U.S. Code 7401 et seq.). In accordance with its responsibility for ensuring high air quality in the Tuxedni Wilderness Area, the Service sought to assess the existing condition of wilderness resources in relation to air quality using the guidelines proposed by Fox et al. (1987). Applying these procedures, the first step in characterizing the plant component is to establish a floristic list that includes for each species distributional information and a commonness rating. Reports on the vascular flora (Tal bot et al. 1995) and lichens (Tal bot et al. 1992) have been published. The bryoflora, in contrast, is virtually unknown. The bryoflora of Alaska is relatively well documented, as summarized by the check­ lists of Frye & Clark (1937- 1947), Worley & Iwatsuki (1970) and Worley (1970). Studies pertinent to Pacific portions of Alaska appeared as early as that of Kurtz (1885) for the Chilkat region. The summary studies of Cardot & Theriot (1902, 1906) also contributed

J Department of Botany, University of British Columbia, Vancouver, British Columbia V6T IZ4, Canada. 2 US. Fish and Wildlife Service, 1011 East Tudor Drive, Anchorage, Alaska 99503, US.A. 3 Biological Science Office, US. Geological Survey, Alaska Science Center, 1011 East Tudor Road, Anchorage, AK 99503, US.A. 92 1. Hattori Bot, Lab, No, 92 2 0 0 2

153'3B'W 37' 36' 35' 34' 153'33'W

60',1'N

10'

Duck 09 ' G} lslc nd

OS'

1 07' N

II LAKE CLARK INLET

NATIONAL PAR K SO'OS'N

SCALE IN KILOMETERS AND PRESERV E

Con tour inter va l = 152 meters

Fig, I, The location and general topography of the Tuxedni Wilderness study area, Alaska, basic information, Evans (1900, 1914) gave scholarly assessments of the hepatics, and Eye­ rdam (1952) provided additional records, Holzinger & Frye (1921) presented the results of the U.S, Kelp Expedition, while Bartram (1938) provided a summary of the mosses known from the Aleutian Islands. Harvill (1948, 1950) made useful assessments of the geography of the mosses of Alaska. Persson (1946-1968), Persson & Shacklette (1969), and Persson & Viereck (1983) have published invaluable reports on scattered incidental collections of bryophytes, many of which were obtained accidentally as mixtures or teased out from vas­ cular plant herbarium specimens. Trelease (1902) and Eyerdam (1955) reported on exten­ sive collections of Sphagna. More recent studies have been made, unfortunately most of them unpublished. Three studies: - the Ph .D. thesis of 1. A. Worley (1972) concerning the bryophytes of the Alexander Archipelago and that of P. G. Davison (1993) concerning the hepatics of the Aleutian Islands, as well as a manuscript produced by D. K. Smith (1984) on the mosses of the same area represent invaluable summaries and syntheses. Peterson et al. (1980) have provided a summary of the mosses of Kodiak Island. Steere (1978) and Steere & Inoue (1978) provide a valuable assessment of the mosses and hepatics of Arctic Alaska. Despite an impressive number of investigations, the bryophytes of the Alaska Peninsu­ la and adjacent islands, including the Tuxedni Wilderness Area, remain inadequately docu­ mented. Access has been, and remains, a major impediment, but we have been fortunate in W B. SCHOFIELD ET AL.: 8ryophytes from Tuxedni Wilderness area, Alaska 93

Fig. 2. Aerial oblique photograph of Chisik Island, Alaska, from the northwest; Duck Island and Cook Inlet are to the left and center and the Tuxedni Channel and the Lake Clark National Park and Preserve are to the right. recent years, largely through support by the U.S. Fish and Wildlife Service, to acquire com­ prehensive collections from many localities in the area. The objectives of this study of Tuxedni Wilderness are to, 1) establish a checklist of the bryophytes, 2) assign a commonness rating - abundant, common, uncommon, and rare - for each species, 3) record habitat information for each species, 4) collect voucher specimens of bryophytes, and 5) compare the bryophyte distribution pattern at different ge­ ographic levels: the northern hemisphere, North America, and Alaska.

GEOGRAPHICAL SETTING Location Tuxedni Wilderness is located at 60 0 08'N, 152°35'W at the head of Tuxedni Bay on the western side of lower Cook Inlet. Chisik, the larger of the two islands, is 10.5 km long and encompasses about 2297 ha, while Duck Island adjacent and east of Chisik is only 0.3 km long and includes 6 ha. Chisik Island is separated from the mainland by Tuxedni Channel which ranges in width from 1.1 to 3.2 km. The topography of Chisik Island is rugged (Wanek 1968). From the southern end of the island the land rises gradually along a ridge to the highest point at 815 m in the north­ ern portion where it drops precipitously along rock cliffs (Fig. 2). Both eastern and western 94 1. Hattori Bot. Lab. No. 92 2 0 0 2

slopes rise steeply from tidal flats to the ridge crest. The study area is found within the "Alaska-Aleutian Province" of the "Alaska Range (Southern Part) Section" of Wahrhaftig (1965) and the "Coastal Zone" of Racine and Young (1978).

Climate Phytogeographically, the Tuxedni Wilderness Area is of interest because it occurs in the northern latitudinal limit of the "maritime zone" (Selkregg 1974: 5, Figure 3). Com­ pared with other climatic zones within Alaska, the maritime zone is distinguished by heavy precipitation, cool summers, and mild winters. Records from the Iniskin climatic station (59°45'N, 153°14 'W), 55 km southwest of Chisik Island on the western side of Cook Inlet, are applicable to the climatic regime for Tuxedni Wilderness (Arctic Environmental Information and Data Center 1989). Mean an­ nual temperature and precipitation recorded for Iniskin between 1954 and 1962 are 0.9°C and 1844 mm, respectively. In contrast, Homer (59°38'N, 151 °33'W), on the eastern side of Cook Inlet, located 80 km southeast of Chisik Island, is classified within the "transition zone" (Selkregg 1974). Homer has a slightly higher mean annual temperature, 2.9°C, and substantially lower pre­ cipitation, 632 mm, for the same reporting period. In addition, total annual snowfall at Homer, 1400 mm, is approximately one third that of Iniskin, 5103 mm. The drier climate of Homer reflects the influence of the Kenai Mountains, with elevations of 1220 to 1830 m. As air is lifted over the mountains, most of its moisture is deposited on the windward side (National Climatic Data Center 1987). Chisik Island receives higher precipitation, because southeasterly winds can penetrate Cook Inlet through an oceanic gap. Based on the ecoclimatic-phytogeographical system of Tuhkanen (1987: 130, Figure 18) for classifying the circumpolar zone, the study area falls within the middle boreal sub­ zone, hyperoceanic (02) sector, and humid (h) province. Using the formula of Hamet-Ahti et al. (1974) to determine climatic subzone, we calculated the mean annual biotemperature for Iniskin to be 3.9°C, placing it within the upper oroboreal subzone, the vertical equiva­ lent of the northern boreal subzone. The elevation of the Iniskin weather station (91 m) probably reflects colder conditions than the basal middle (oro-) boreal subzone at sea level.

Geology and Soils Tuxedni Wilderness is underlain by highly fossiliferous sedimentary bedrock, Middle­ to-Upper Jurassic, (Detterman and Hartsock 1966), including siltstone, sandstone, and con­ glomerate. Surficial deposits occur on Chisik Island along the west-central shore (an allu­ vial fan) and on the northern end (elevated beach ridges). Soil development occurs on undifferentiated alluvium and coarse rubbly deposits on steep mountains and hills (Karlstrom et al. 1964). In a reconnaissance survey of the soils of Alaska, Rieger et al. (1979) mapped Tuxedni Wilderness Area within the "Typic Cryan­ depts, very gravelly, hilly to steep - Rough Mountainous Land Association" within the "Alaska Peninsula and Southwestern Islands Land Resource Area." According to Rieger et al. (1979), the dominant soils under 600 m elevation are Typic Cryandepts. These soils are covered with a mat of litter and consist of strongly acid, dark reddish brown volcanic ash W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 95

Fig. 3. Interior view of an Alnus viridis subsp. sinuata thicket with an understory of Oplopanax horridus on an alluvial fan in northwestern Chisik Island. On the floor of such habitats occur Brachythecium refiexum, Bryhnia hultenii, Plagiothecium denticulatum , and Plagiothecium laetum; epiphytes on Alnus viridis subsp. sinuata include Dicranum tauricum, Iwatsukiella leucotricha, Pseudoleskea stenophylla , and Vlota drummondii.

(25- 50 cm) over dark brown gravelly loam.

Vegetation The small-scale vegetation map of Alaska (Joint Federal-State Land Use Planning Commission for Alaska 1973) classifies the vegetation of Tuxedni Wilderness Area as dominated by Alnus viridis subsp. sinuata thickets, with an understory of grasses and ferns. In a large-scale (1 : 63,360) vegetation map of Tuxedni Wilderness, Racine and Young (1978) mapped six types: (1) coastal Sitka spruce forest, (2) alder shrub thicket, (3) dry alpine tundra, (4) alpine barrens, (5) Calamagrostis grassland, and (6) Leymus shoreline. General descriptions of the plant communities are as follows (Talbot et al. 1995) and vascular plant nomenclature follows USDA, NRCS (1999). Broadleaf deciduous thickets of Alnus viridis subsp. sinuata predominate on well-drained sites at lower to middle elevations (Fig. 3, Fig. 4). Typical understory species in the thickets are the shrubs, Oplopanax hor­ ridus, Rubus spectabilis, and Sambucus racemosa; forbs, Athyriumfilix-femina, Dryopteris expansa, Gymnocarpium dryopteris, Trientalis europaea, and Veratrum viride; and graminoid, Calamagrostis canadensis. The Alnus viridis subsp. sinuata thickets of Tuxedni 96 1. Hattori Bot. Lab. No. 92 2 002

Fig. 4. General aspect of an Empelrum nigrum - Cassiope stelleriana hummocky heath looking toward the southern end of Chisik Island, Al aska. Characteristic bryophytes of these habitats include Andreaea a/peslris, Barbilophoziajloerkei, Echinophyllum sachalinen­ sis, Hypnum plicatulum, and Pseudoleskea baileyi.

Wilderness correspond to the "Subalpine Zone of glaciated uplands and moist mountain valleys .. . (with) . . . a mosaic of Alnus crispa thickets and tall-growing grass and forb com­ munities," described by Mitchell (1968, p. 45) for south-central Alaska, which includes the Tuxedni area. Similarly, Hamet-Ahti (1976, p. 56, fig . 5) characterized an upper oroboreal subzone, approximately equivalent to subalpine, by the dominance of Alnus viridis subsp. sinuata for the oceanic boreal zone of western North America. Other scrub types are less abundant and include Rubus spectabilis thickets on well-drained lower slopes, Salix pul­ chra thickets on protected, well-drained upper elevation sites, and Salix barclayi swamps. Dwarf shrub communities achieve dominance above about 610 m. The most extensive community types are Empetrum nigrum - Cassiope stelleriana hummocky heath (Fig. 5) on protected, moderately well-drained sites, Vaccinium uliginosum - Empetrum nigrum heath on slightly more exposed sites, and Arctostaphylos alpina - Cladina rangiferina heath on the most wind-exposed sites. Late-melt snowbed, Luetkea pectinata scarcely vegetated communities, are associated with the Empetrum - Cassiope hummocky heaths. Herbaceous vegetation occurs primarily as tall forb meadow communities at middle elevations in a mosaic with Alnus viridis subsp. sinuata thickets (Fig. 5). Meadows are characterized by the forbs, Athyriumfilix-femina, Chamerion angustifolium, Senecio trian- W. B. SCHOFI ELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 97

Fig. 5. Mid-elevation (550 m) mosaic of Alnus viridis subsp. sinuata thickets with an understory dominated by the ferns Athyrium filix-femina and Dryoperis expansa, and tall forb meadows characterized by the forbs Chamerion angustifolium, Senecio triangularis, and Veratrum viride. In such meadow habitats the following can be found: Barbilophoziafloerkei, Dicranum scoparium, Hypnum lindbergii, Pleurozium schreberi, Polytrichum commune, and Tortula norvegica. gularis, and Veratrum viride, and the graminoid, Calamagrostis canadensis. Other herba­ ceous vegetation includes Calamagrostis canadensis - Sanguisorba stipulata meadows along stream lets, Chamerion angustifolium - Lupinus nootkatensis meadows on beach ridge deposits, Leymus mollis graminoid meadows and Honckenya peploides forb meadows along beach shores, and hanging garden communities on steep cliff walls with the forbs, Artemisia tilesii, Chamerion latifolium, Mimulus guttatus, and Rhodiola rosea, and the graminoids, Hordeum brachyantherum and Poa macrocalyx. Mire communities occur as relatively isolated types in poorly drained depressions. They are usually dominated by Trichophorum caespitosum. Characteristic mire species in­ clude the dwarf shrubs, Andromeda polifolia, Betula nana, and Oxycoccus microcarpus; forbs, Comarum palustre, Drosera rotundifolia, Erigeron peregrinus, Gentiana dou­ glasiana, Platanthera dilatata, Spiranthes romanzoJfiana; and the graminoids, Eriophorum angustifolium and Iris setosa. Forest communities are minor components of the vegetation. Picea sitchensis forest occurs on the northernmost portion of Chisik Island on well-drained sites and as isolated individuals or stands. Populus balsamifera subsp. trichocarpa forest communities are as so- 98 1. Hattori Bot. Lab. No. 92 2 0 0 2

ciated with well-drained glacial outwash.

Post-glacial Vegetation In a study of the post-glacial history of the Cook Inlet region, Ager et al. (1985) demonstrated that deglaciation started at 14,000yr B.P. but may have lagged in valleys until as late as 10,500 yr B.P. Basal pollen assemblages suggest the initial vegetation types con­ sisted of herb-shrub tundra or shrub-herb tundra dominated by Betula cf. nana, Salix, and Ericales. At an interval about 10,500- 9,000 yr B.P., the tundra was replaced by Populus woodlands interspersed with Salix thickets and shrub tundra. Alnus subsequently migrated into the area about 9,500 yr B.P. Boreal spruce (Picea glauca, P mariana) incursions through the Matanuska Valley began about 8,500- 8,200 yr B.P. and expanded over much of the region by mid-Holocene. Forests of Picea sitchensis and Tsuga mertensiana that presently occupy coastal areas and mountain valleys developed relatively recently, within the past 2,000 years.

METHODS Prior to initiating field studies, a marine circumnavigation was conducted in April 1987 to gain familiarity with the study area. Following this reconnaissance, color-infrared aerial photographs (scale I : 60,00) were interpreted in the laboratory to delineate habitat types and identify potential sampling locations selected to represent the environmental variation within the area. Field studies began in 1987, continued during the summer of 1988, and completed in June 1993. The 1987 field schedule was divided into two periods in order to collect both early- and late-flowering plants, and fruiting bryophytes: 26 June to II July and 24 August to 5 September. Persistent snow cover at upper elevations restricted plant collecting to lower elevations during the early period. Field studies were continued in 1988 (26 August to 15 September) and 1993 (10 to 24 June) to insure that all significant sites were sampled. Three hazardous sites were not well­ explored for safety reasons; these were talus slopes, steep rock cliffs, and the northeastern coast. Despite these exceptions we feel the list of species is nearly complete and further studies would add only a few species. All species included are documented by specimens at the University of British Colum­ bia Herbarium and replicates of many are deposited at the herbarium of the University of Alaska at Fairbanks. Additional replicates, when available, are deposited at Duke Universi­ ty Herbarium, Missouri Botanical Garden Herbarium and at the herbarium of New York Botanical Garden as well as other world herbaria. In collecting, an attempt was made to in­ clude a representative spectrum of habitat types over complex topographic gradients. Bryophyte nomenclature follows Anderson (1990) for Sphagnum, Anderson et al. (1990) for mosses, with some exceptions, and Stotler & Crandall-Stotler (1977) for liver­ worts. Vascular plant nomenclature follows USDA, NRCS (1999). A large part of the col­ lection was made and determined by WBS. Other collections were made by Stephen Talbot and Sandra Looman Talbot; most of these bryophytes were determined by WBS, Sphag­ num by Richard E. Andrus, and some bryophytes by William R. Buck. W. B. SCHOFIELD ET AL. : Bryophytes from Tuxedni Wilderness area, Alaska 99

CHECKLIST FORMAT Each species is followed by a series of categories: habitat, elevational position in the landscape, fruiting condition is indicated as sterile or cfr. (= sporophytes are present in some plants), collection number(s), commonness rating (in parentheses; see below), and geographical distribution. Commonness rating classes follow Duncan and Meacham (1986): (1) abundant = very likely to be encountered; nearly always found in appropriate habitats, sometimes forming dense stands; (2) common = likely to be encountered; (3) uncommon = unlikely to be encountered and sometimes not present in appropriate habitats; and (4) rare = extremely unlikely to be encountered, often not present in appropriate habitats, and often restricted to a small number of sites. The cited specimen numbers 98666-99217 are those of Schofield. All others are those of Talbot and Looman Talbot. The following list is conservative. There remain some specimens that are undeter­ mined. Most of these belong to taxonomically poorly understood genera or represent taxa that were available only as vegetative material where sporophytes are critical to accurate determination. A concentrated attempt was made to acquire sporophyte-bearing material, but many taxa either lack sporophytes in island populations, or sporophytes were infre­ quent and not discovered in spite of a concentrated attempt. Further study of the material may make determinations possible, but the current literature is insufficient to do so.

ANNOTATED CATALOG OF BRYOPHYTES Taxa in the checklist are arranged in alphabetical order.

Mosses (230 taxa) Abietinella abietina (Hedw.) Fleisch., exposed outcrops, sea-level and subalpine; sterile; 98674, 98803,99210. (4) Circumboreal. Amblystegium serpens (Hedw.) Schimp., shaded rock, soil, logs, and tree bases, lower elevations; cfr. ; 98672, 98884, 98958, 98959. (3) Cosmopolitan. Amphidium lapponicum (Hedw.) Schimp., shaded cliffs, lower elevations; cfr.; 98669, 98761A, 98864,98908, 87-153, 87-552, 87-68, 87-71,87-76, 87-77, 87-78A, 87-83. (2) Circumboreal. A. mougeotii (B. S. G.) Schimp., shaded or open cliffs, lower elevations; sterile; 99039, 99045, 99054. (3) Boreal-Interrupted. Andreaea alpestris (Thed.) Schimp., terrestrial, tundra, subalpine; sterile; 99102, 99105. (3) Mon­ tane-Wide Disjunct. A. blyttii Schimp., on rock, late snow melt sites, subalpine; cfr. ; 99121, 99125, 88-11-14, 88-15-15. (3) Circummontane. A. nivalis Hook., rock, late snow-bed sites, subalpine; sterile; 99120, 88-1208A . (2) Circummontane. A. rupestris Hedw., on rock at all elevations; cfr. ; 98682, 98695, 98762, 98801, 98815, 98823, 98825, 98831,99106, 99171 , 99184, 87-51,87-877,87-426. (1) Circumboreal. Anoectangium aestivum (Hedw.) Mitt., humid shaded cliff, lower elevations; sterile; 87-138. (4) Cir­ cumtemperate. 100 J. Hattori Bot. Lab. No. 92 2 0 0 2

Map I (left). North American distribution of Antitrichia californica Sull. Map 2 (right). North American distribution of Brachydontium olympicum (Britt.) McIntosh & Spence.

Anomobryum filiforme (Dicks.) Solms, damp shaded cliff near sea-level; sterile; 98671, 98903, 99202. (3) Montane-Wide Disjunct. Antitrichia californica Sull., open cliff, near sea level; sterile; 98876. (4) Temperate-Western North ArnericalEurope Disjunct. (Map I) A. curtipendula (Hedw.) Brid., cottonwood trunk, near sea level; sterile; 99008. (4) Temperate-Inter­ rupted. Arctoafulvella (Dicks.) Bruch & Schimp., cliff crevices and outcrops, subalpine; cfr.; 98817, 99119, 87-943,88-11-10,88-11-12, 88-12-10,88-13-9. (4) Circummontane. Aulacomnium palustre (Hedw.) Schwaegr., mires and heath slopes, various elevations; sterile; cfr.; 98828,88-47-18,87-24-17. (I) Cosmopolitan. A. turgidum (Wahlenb.) Schwaegr., sea stack near sea level; sterile; 98893. (4) Arctic-Alpine. Bartramia ithyphylla Brid., cliff crevices, various elevations; cfr.; 98715, 98864A , 99157, 87-791, 87- 699. (2) Arctic-Alpine. B. pomiformis Hedw., cliff ledges and bases, various elevations; cfr.; 98874, 99056, 87-518A, 87-86, 87-515. (3) Circumboreal. Bartramiopsis lescurii (James) Kindb., soil over outcrop, stream canyon, lower elevations; cfr.; 99057, 87-542A, 87-525. (3) Boreal-AmphiPacific. Blindia acuta (Hedw.), Bruch & Schimp., damp cliffs and stream margins, lower elevations; cfr.; 98922,87-157,87-159,87-160,87-162,87-33, 87-91 , 87-92, 87-96, 87-103, 87-607. (2) Bore­ ai-Interrupted. W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 101

Brachydontium olympicum (Britt.) McIntosh & Spence, exposed conglomerate outcrop, subalpine; cfr. ; 98746. (4) Montane-AmphiPacific. (Map 2) Brachythecium albicans (Hedw.) Schimp., open well-drained sites at all elevations; sterile; 98708, 98858,88-1209,88-1401,87-6-9,87-13-20,87-18-9, 87-133, 87-409, 87-428A, 1420F. (I) Cir­ cumboreal. B. erythrorrhizon B. S. G., epiphytic on alder and cottonwood; sterile; 87-4-6, 87-8-12, 87-16-18, 87- 40-14, 87-40-22. (2) Boreal-Interrupted. B. holzingeri (Grout) Grout, in cottonwood stand, lower elevations; sterile; 87-16-18. (4) Montane­ Western North America Endemic. B. populeum (Hedw.) Schimp., cottonwood stand, lower elevations; sterile; 87-40-11. (4) Circumbo­ real. B. plumosum (Hedw.) Schimp., shaded cliff and boulders, lower elevations; c/r.; 98875, 99012. (2) Cosmopolitan. B. refiexum (Starke) Schimp., epiphytic, on logs and terrestrial, all elevations; cfr.; 98931, 98944, 98945, 98848, 98970, 98972, 98994, 99143, 99158, 99169, 88063-X-19, 87-1-10, 87-4-5, 87- 43-7,87-12-7,87-35-9,87-39-5,87-45-4,87-48-8. (I) Circumboreal. B. rivulare Schimp., wet cliff and swamps, lower elevations; sterile; 98912, 98953, 98962, 561, 604. (3) Circumboreal. B. salebrosum (Web. & Mohr) Schimp., epiphytic and decaying logs, lower elevations; sterile; 99001 , 87-3-8,87-12-9. (3) Circumboreal. B. starkei (Brid.) Schimp., reclining trunks of alder; cfr.; 98675, 98717, 98722, 98946, 98968, 98971. (2) Circumboreal. B. turgidum (Hartm.) Kindb., cliff face, subalpine; sterile; 87-161. (4) Arctic-Alpine. B. velutinum (Hedw.) Schimp., soil bank, lower elevation; cfr.; 98947. (4) Circumboreal. Bryhnia hultenii Bartr., shaded cliff bases, stream margins, and alder thickets, all elevations; sterile; 98865,99032,99076,582. (2) Boreal-AmphiPacific. Bryoerythrophyllum inaequalifolium (Tayl.) Zand., seepy cliff near sea level; sterile; 98907. (4) Mon­ tane-Wide Disjunct. B. recurvirostre (Hedw.) Chen, cliff faces, near sea level; cfr.; 87-135, 87-136, 87-599, 87-601, 88- 1420J. (3) Cosmopolitan. Bryum argenteum Hedw., earth bank near gull roost, sedge mire, lower elevations; sterile; 98854, 87- 32-4. (4) Cosmopolitan. B. caespiticium Hedw. , cliff crevices, near sea level; cfr.; 576, 99144. (3) Cosmopolitan. B. capillare Hedw. , upper stony beach near sea level; sterile; 1402. (4) Cosmopolitan. B. creberrimum Tayl., streamside and in cottonwood stand, low elevations; cfr.; 1298, 1362A. (3) Cos- mopolitan. B.fiaccidum Brid., cliff seep, near sea level; sterile (gemmae); 595. (4) Boreal-Interrupted. B. miniatum Lesq., cliff shelf, near sea level; sterile; 99148. (4) Temperate-Interrupted. (Map 3) B. turbinatum (Hedw.) Turn., conglomerate cliffs, near sea level; cfr.; 98883. (3) Cosmoplitan. B. weigelii Spreng., seepage, near snowbeds, subalpine; sterile; 98702, 98736, 99087, 87-877. (3) Circumboreal. Calliergon cordifolium (Hedw.) Lindb., brook in alder swamp, lower elevation; sterile; 98954, 98955, 98956. (4) Circumboreal. C. megalophyllum Mik., fen margin, lower elevations; sterile; 99131. (4) Boreal-Interrupted. (Map 4) C. stramineum (Brid.) Kindb., in Sphagnum at edge of fen and in snowbed areas, subalpine and mid­ elevations; sterile; 99061,99086,99090,99108. (2) Circumboreal. 102 J. Hattori Bot. Lab. No. 92 2 002

Map 3. North American distribution of Bryum miniatum Lesq.

Map 4. North American dis­ tribution of Calliergon megalo­ phyllum Mik. W. B. SCHOFI ELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 103

Campylium hispidulum (Brid.) Mitt., mineral soil bank, low elevations; cfr.; 98943. (4) Temperate­ Wide. C. stellatum (Hedw.) C. Jens., damp areas in wetland, mid-elevations; sterile; 99079, 99109, 88-47- 23. (2) Circumboreal. Ceratodonpurpureus (Hedw.) Brid., open sites, all elevations; cfr.; 98742, 98846, 99138, 87-223, 87- 9-22,87-412. (2) Cosmopolitan. Cinclidium subrotundum Lindb. near pond and streamlet margins in fen, mid-elevations; cfr.; 99097, 99129. (3) Arctic-Circumpolar. Cirriphyllum cirrosum (Schwaegr.) Grout, seepy or dry cliffs, near sea level; sterile; 98911, 99047, 99145. (4) Arctic-Alpine. Claopodium bolanderi Best, rock outcrop, subalpine; sterile; 87-693. (4) Temperate-Western North America Endemic. Climacium dendroides (Hedw.) Web. & Mohr, moist and dry sites near cliff bases and water-courses, various elevations; sterile; 98759, 99050, 99175, 87-606, 87-2-16. (2) Circumboreal. Conostomum tetragonum (Hedw.) Lindb., open slope, subalpine; cfr.; 98692. (4) Arctic-Alpine. Cratoneuron filicinum (Hedw.) Spruce, damp cliffs and swamps, lower elevations; sterile; 98916, · 98957, 99082,87-612. (3) Circumboreal. Cynodontiumjenneri (Schimp.) Stirt., cliffs , subalpine and sea level; cfr.; 98673, 88-4-37. (4) Boreal­ Interrupted. C. strumiferum (Hedw.) Lindb., lichen heath, subalpine; cfr.; 88-5-33, 88-34-39. (4) Circumboreal. Desmatodon latifolius (Hedw.) Brid., soil of tundra, subalpine; cfr.; 98711, 98754A . (4) Arctic­ Alpine. Dichodontium olympicum Ren. & Card., soil on heath slope, subalpine; sterile; 98793. (4) Montane­ Western North America Endemic. (Map 5) D. pellucidum (Hedw.) Schimp., damp areas on cliffs and near streams, lower elevations; sterile; 99035,87-613. (2) Boreal-Interrupted. Dicranella grevilleana (Brid.) Schimp., earth bank on cliff, lower elevation; cfr.; 98885. (4) Circum­ boreal. D. heteromalla (Hedw.) Schimp., earth under alders, mid and low elevations; sterile; 98769A , 98839. (4) Circumboreal. D. palustris (Dicks.) Crundw., seepage sites, subalpine; sterile; 98737, 87-281,87-605. (3) Boreal-In­ terrupted .. Dicranoweisia crispula (Hedw.) Lindb., on cliffs and boulders, all elevations; cfr.; 98703, 99042, 99116,87-48,87-50,87-60, 87-571A, 87-419, 87-436, 87-437A. (1) Arctic-Alpine. Dicranum acutifolium (Lindb. & AmelI) C. Jens. ex Weinm., lichen heath, subalpine; sterile; 99093, 88-5-30,88-24-11,88-66-30,718,726,732,740,818. (2) Circumboreal. D. elongatum Schwaegr., lichen tundra, subalpine; sterile; 88-5-21, 746A . (4) Circumboreal. D. fuscescens Turn., rotten wood and tree bases, lower elevations; cfr. ; 98813, 98949, 98992. (3) Cir­ cumboreal. D. majus Srn., in Sitka spruce stand, dwarf shrub heath, and mires, at all elevations; cfr.; 98980, 88- 1290£1,87-2-20, 87-2-22,87-467. (3) Boreal-Interrupted. D. muehlenbeckii B.S.G., crowberry heath, subalpine; sterile; 88-73-10. (4) Circumboreal. D. scoparium Hedw., heath, fens, snowfield meadow, Sitka spruce forest, cfr.; 98699, 98772, 98775, 98820,98998,99058, 99059, 99063,98684,98800, 98977, 88-1291, 87-456. (2) Circumboreal. D. tauricum Sapeh., epiphytic on Sitka spruce and alder, lower elevations and subalpine; sterile; 98981, 99072, 991 85, 87-112, 87-233, 88-1374H. (3) Temperate-Western North AmericalEu- 104 J. Hattori Bot. Lab. No. 92 2 0 0 2

Map 5 (left). World distribution of Dichodontium o/ympicum. Ren. & Card. Map 6 (right). North American distribution of Echinophyllum sachalinensis (Lindb.) O'Brien in O'Brien & Horton.

rope Disjunct. D. undulatum Brid., edge offen, mid-elevations; cfr.; 99062, 99075,88-33-13. (3) Circumboreal. Didymodon rigidulus Hedw. , cliff seep, near sea level; sterile (gemmae); 605. (4) Circumboreal. D. vinealis (Brid.) Zand. var.jl.accidus (Bruch & Schimp.) Zand., shaded cliff, lower elevations; ster­ ile; 98888, 87-587, 598, 600. (3) Circumtemperate. Distichium capillaceum (Hedw.) Bruch & Schimp., shaded cliff, lower elevations; cfr.; 98909, 87- 507, 87-524,51 7. (3) Cosmopolitan. Ditrichum heteromallum (Hedw.) Britt., conglomerate cliff and earth banks, lower elevations; cfr.; 98883A, 99029, 99164. (4) Temperate-Western North America/Europe Disjunct. D. zonatum (Brid.) Kindb., damp cliff near snowbed, subalpine; sterile; 99114. (4) Temperate-West­ ern North America/Europe Disjunct. Drepanocladus aduncus (Hedw.) Warnst. , edge of snowbed; sterile; 99085. (3) Cosmopolitan. D. polycarpus (Voit.) Warnst., aquatic in wetland bluejoint meadow, mid-elevation; sterile; 98952. (3) Circumboreal. Encalypta ciliata Hedw., shaded cliff, lower elevation; cfr.; 98868., 98872. (4) Circumboreal. E. rhaptocarpa Schwaegr., conglomerate cliff shelf, sea level; cfr.; 98666. (4) Circumboreal. Echinophyllum sachalinensis (Lindb.) O'Brien in O'Brien & Horton, depressions in crowberry tun- dra, subalpine; sterile; 98826, 98802, 98833; 88-22-9. (4) Montane-AmphiPacific. (Map 6) Eurhynchium pulchellum (Hedw.) Jenn., shaded cliff and soil, lower elevation; sterile; 98847A , 98873, 98942. (3) Circumboreal. Fissidens bryoides Hedw., shaded cliff, stream let canyon, lower elevation; cfr. ; 99154. (4) Circum- W. B. SCHOFIELD ET AL.: 8ryophytes from Tuxedni Wilderness area, Alaska 105

temperate. Funaria hygrometrica Hedw., maritime cliff, near sea level; cfr. ; 98915, 98921, 1420K. (3) Cos­ mopolitan. Grimmia donniana Srn., on boulders in tundra and near sea level, cfr. ; 98852, 98897, 87-408, 87-414, 87-423,87-431, 760, 769. (3) Circummontane. G. tenerrima Ren. & Card., cliff top, subalpine; cfr.; 98706. (4) Circummontane. G. torquata Hornsch., dry outcrop faces near sea level; sterile; 99139, 87-514. (4) Boreal-Interrupted. Gymnostomum aeruginusum Srn., seepy cliff, near sea level; cfr.; 98902. (4) Circumboreal. Hamatocaulis vernicosus (Mitt.) Hedenas, edge of fen streamlet, mid-elevation; sterile; 99132. (4) Circumboreal. Herzogiella adscendens (Lindb.) Iwats. & Schof., shaded seaside cliff near sea level; sterile; 98847. (4) Boreal-AmphiPacific. H. striatella (Brid.) Iwats., shaded cliff ledges and humus banks in alder thickets, lower elevations; cfr.; 99055, 99186, 88-63-12. (4) Boreal-Interrupted. Hygrohypnum alpinum (Lindb.) Loeske, rock in streamlet, low elevation; cfr. ; 99025. (4) Circum­ montane. H. bestii (Ren. & Bryhn) Broth., on rock of wet cliff and in stream, low elevations; sterile; 99168, 98913,87-342,87-543. (3) Boreal-AmphiPacific. H. molle (Hedw.) Loeske, on wet rock, near snowbed and splash zone of waterfall, subalpine and low elevation; sterile; 611 , 613,914. (3) Circummontane. H. ochraceum (Turn.) Loeske, rock and stick in stream, low elevations; sterile; 99033, 87-30, 87-545, 87-614, 88-1295H. (3) Boreal-Interrupted. Hylocomiastrum pyrenaicum (Spruce) Fleisch., depression in mountain slope, mire slope, mid-eleva­ tions; sterile; 88-18-13, 88-21-20, 88-32-20. (4) Circurnboreal. Hylocomium splendens (Hedw.) Schimp., terrace of escarpment, subalpine; sterile; 98709, 87-285. (2) Circumboreal. Hymenostylium insigne (Dix.) Podp., damp shaded cliff, near sea level; sterile; 99204, 98910, 87-158, 87-517A, 579, 596. (3) Temperate-Western North AmericaJEurope Disjunct. (Map 7) H. recurvirostre (Hedw.) Dix., damp cliff crevice, near sea level; sterile; 98919. (4) Circumboreal. Hypnum callichroum Funck, damp bank of steamlet, low elevations; sterile; 98778. (4) Circumboreal. H. cupressiforme Hedw. var subjulaceum Mol., cliff shelves, near sea level; sterile; 98855, 98856, 98877,99077,99216,87-84,87-85. (4) Temperate-Interrupted .. H. lindbergii Mitt., damp grassy slopes, depressions in crowberry heath, and peaty pond edges, main­ ly subalpine; sterile; 98731,98835,99088. (3) Circumboreal. H. plicatulum (Lindb.) Jaegr., heath and dwarf shrub tundra, subalpine; sterile; 98794, 98819, 87- 733,87-734,88-3-32,88-5-31, 88-34-28,88-5-31, 88-10-20. (3) Boreal-Interrupted. H. revolutum (Mitt.) Lindb., dry boulders, boulder slope, near sea level; sterile; 98899, 99209. (4) Circumboreal. Isopterygiopsis pulchella (Hedw.) Iwats., shaded cliff crevice, rocks, and epiphytic on cottonwood base, low elevations; cfr. ; 99043, 87-40-21, 88064-7. (4) Circumboreal. Isothecium stoloniferum Brid., twig of fallen Sitka spruce, low elevations; sterile; 98987. (4) Temper­ ate-Western North America Endemic. Iwatsukiella leucotricha (Mitt.) Buck & Crum, trunks of cottonwood, Sitka spruce, and alder; cfr. ; 98840, 98995, 88-1375L, 87-13-9. (4) Boreal-AmphiPacific. (Map 8) Kiaeria blyttii (Schimp.) Broth,. heath near snowbed, subalpine; cfr. ; 98732, 88-4-27, 88-7-32, 87- 425, 88014-13, 792, 848, 849A. (3) Circummontane. 106 1. Hattori Bot. Lab. No. 92 2 0 0 2

Map 7 (left). North American distribution of Hymenostylium insigne (Dix.) Podp. Map 8 (right). North American distribution of 1watsukiella leucotricha (Mitt.) Buck &Crum.

K.falcata (Hedw.) Hag., snowbed and clifftop, subalpine; cfr. ; 98704, 88-13-7. (3) Circummontane. Leptobryum pyriforme (Hedw.) Wils., soil by building, near sea level; cfr.; 98895. (4) Cosmopolitan. Lescurea saxicola (Schimp. in B. S. G.) Milde, stabilized shaded boulders on slope, mid-elevations; sterile; 99197. (4) Circummontane. Leskeella nervosa (Brid.) Loeske, horizontal rock face and cliff, near sea level; sterile; 98849, 98851, 98929, 99137, 691. (4) Circumboreal. Limprichtia revolvens (Sw.) Loeske, wet depression in fen and snowbed seepage, subalpine; sterile; 99095, 98723. (3) Circumboreal. Mielichhoferia macrocarpa (Hook.) Bruch & Schimp., wet cliff crevices, low elevations; cfr. ; 99133, 577, 593. (4) Temperate-AmphiPacific. Mnium spinulosum Bruch & Schimp., rotten log in cottonwood stand, low elevations; cfr.; 98969. (4) Boreal-Interrupted. M. thomsonii Schimp., humid cliff face, low elevations; sterile; 87-79, 87-517, 87-522. (4) Circumbo­ real. Myurellajulacea (Schwaegr.) Schimp., seepy cliff, near sea level; sterile; 98900. (4) Circumboreal. Oedipodium griffithianum (Dicks.) Schwaegr., damp shaded cliff, near sea level; cfr.; 98670, 87-39, 87-147, 87-148,87-149,87-150. (4) Circummontane. Oligotrichum hercynicum (Hedw.) Lam. & DC., mineral soil in heath and near snow patches, sub­ alpine; cfr. ; 98693, 98777, 99064, 88-12-6, 88-2-6, 87-1011 . (3) Circummontane. 0. parallelum (Mitt.) Kindb., edge of snowbed and stream margins, subalpine; sterile; 98734, 98735, W. B. SCHOFIELD ET AL. : Bryophytes from Tuxedni Wilderness area, Alaska 107

98766, 88-1295G. (4) Boreal-AmphiPacific. Oneophorus wahlenbergii Brid., trunk of Sitka spruce, lower elevations; efr.; 98983. (4) Circumboreal. Orthotriehum anomalum Hedw. , cliffs, alder stumps and boulders, near sea level; efr. ; 98845, 98881, 99147, 697. (3) Circumtemperate. 0. obtusifolium Brid., on trunk of cotton wood, near sea level; efr. ; 98997. (4) Circumboreal. 0. pal/ens Bruch, trunks of elderberry and mountain ash, lower elevations; efr. ; 98887, 87-1-21. (4) Boreal-Interrupted. 0. pulehellum Brunt., alder trunks, lower elevations; efr. ; 98882, 99000. (4) Temperate-Western North America/Europe Disjunct. 0. speeiosum Nees, epiphytic on alder and cottonwood; eft: ; 99007, 99011, 87-128, 87-443. (3) Cir­ cumboreal. 0. striatum Hedw., epiphytic on alder, paper birch and cottonwood, lower elevations; efr.; 98926, 98927, 98941,99002, 87-12-5,87-14-7,87-40-28, 87-450. (3) Temperate-Interrupted. Oxystegus tenuirostris (Hook. & Tayl.) A. 1. E. Srn., cliffs by streamlet, lower elevations; sterile; 98784,99156,571. (4) Circumtemperate. Paludella squarrosa (Hedw.) Brid., rare in fen, subalpine; sterile; 99096, 88-45-11 . (4) Circumboreal. Philonotis eapillaris Lindb., mineral soil, lower elevations; efr.; 99153, 87-32-11. (4) Temperate-In­ terrupted. P. fontana (Hedw.) Brid., damp cliffs and wetlands, lower and middle elevations; efr.; 98923, 88- 1290E2, 87-101, 87-440. (2) Circumboreal. Plagiobryum zierii (Hedw.) Lindb., damp cliff, near sea level; sterile; 99044. (4) Circummontane. Plagiomnium affine (Bland.) T. Kop., earth in cottonwood stands, swampy areas, lower elevations; cfr. ; 98925; 98930, 98960, 98967. (3) Circumboreal. P. cuspidatum (Hedw.) T. Kop., epiphytic on alder, lower elevations; cfr.; 107, 116. (4) Circumboreal. P. insigne (Mitt.) T. Kop ., moist sites, lower elevations; sterile; 87-603, 566. (3) Temperate-Western North America Endemic. P. medium (Bruch & Schimp.) T. Kop. , alder swamps and cottonwood stands, lower elevations; efr.; 98961 , 98963,87-3-5, 87-16-10, 87-16-19A, 87-19-15. (2) Circumboreal. Plagiopus oederiana (Srn.) Crum & Anderson, moist rocks in stream canyon, lower elevations; ster­ ile; 87-526, 763B, 768A . (4) Circumboreal. Plagiothecium eavifolium (Brid.) Iwats., soil over rocks in shaded alder thickets and shaded cliffs, lower elevations; sterile; 98757, 98812, 99135, 87-141, 87-163, 87-509, 87-94. (2) Temperate­ Interrupted. P. dentieulatum (Hedw.) Schimp., floor of alder thickets, depressions on slopes, subalpine and lower elevations; cfr. ; 99180, 88-20-10, 87-4-4, 87-35-10, 87-37-12, 87-39-6, 88-63-11, 88-20-10, 87- 98. (1) Circumboreal. P. laetum Schimp., humus, rotten wood in wooded or shrubby sites, occasionally on rock, at all eleva­ tions; efr. ; 98720, 98859, 87-21-19, 87-47-7, 87-4-7, 88-24-12, 88-71-17, 87-6-10, 87-66, 87- 139A, 87-140, 87-508. (1) Boreal-Interrupted. P. undulatum (Hedw.) Schimp. in B. S. G., on peat moss in Sitka spruce forest; lower elevation; ster­ ile; 454. (4) Temperate-Interrupted. (Europe-W. NA). Pleurozium schreberi (Brid.) Mitt., heath, boulder slopes, and Sitka spruce forest floor, all elevations; sterile; 98696, 98716, 99208, 87-229, 87-725. (2) Circumboreal. Pogonatum dentatum (Brid.) Brid., base of earth slide, lower elevation; sterile; 99166. (4) Circumbo­ real. P. urnigerum (Hedw.) P. Beauv., mineral soil in heath and at cliff bases, subalpine and at lower eleva- 108 J. Hattori Bot. Lab. No. 92 200 2

tions; cfr.; 98796, 98878, 98892, 88-38-14. (3) Circumboreal. Pohlia cruda (Hedw.) Lindb., mineral soil banks, at all elevations; cfr.; 98740, 87-31, 87-422, 87-429. (2) Circumboreal. P drummondii (c. Mull.) Andrews, damp mineral soil bank, lower elevations; cfr.; 98978. (4) Cir­ cummontane. P elongata Hedw., clayey bank by cliff, lower elevations; cfr.; 98973, 193. (4) Boreal-Interrupted. P nutans (Hedw.) Lindb., soil by building, near sea level ; cfr.; 98896, 87-379, 87-380, 87-429, T3222-12. (4) Circumboreal. P proligera (Kindb.) Lindb., mineral soil banks, lower elevations; sterile (gemmae); 98976, 98739, 99014,99015. (I) Circumboreal. P sphagnicola (Bruch & Schimp.) Lindb. & Ameli, Sphagnum in fen, mid-elevations; sterile; 99067. (4) Circumboreal. P wahlenbergii (Web. & Mohr) Andrews, seepage area, edge of pond, lower elevation; sterile; 99089, 872,902. (3) Cosmopolitan. Polytrichastrum alpinum (Hedw.) G. L. Srn., on overturned roots, cliff shelves, tundra, shrubby or wooded sites, all elevations; cfr.; 98986, 87-132, 87-142, 87-143, 87-20-33, 88-13 73A, 87-53 , 87-294,87-796, 87-763A, 88-12951, 88-17-17, 88-18-1 9, 88-21-15, 88-71-16, 88-18-19, 88-21- 15, 87-16-20, 87-523, 88-2 7-16, 88-74-14, 87-53, 87-695, 87-697, 87-763A , 87-294, 87-427, 87-944. (1) Circumboreal. Polytrichum commune Hedw. , heath tundra, forb meadows, higher elevations; sterile; 98795, 99068, 88-19-8,88-25-9,88-70-24. (3) Cosmopolitan. Pformosum Hedw., humus bank and upper stony beach, near sea level; sterile; 99183, 1400. (4) Cir­ cumboreal. P juniperinum Hedw. , heath tundra, low shrub thickets, shaded humus, all elevations; sterile; 88-4- 32,88-20-9,88-68- 15,99188. (2) Cosmopolitan. P piliferum Hedw., exposed acid mineral soil, at all elevations; sterile; 87-304, 87-418, 87-944, 88- 1208,88-8-31 ,98687. (3) Cosmopolitan. P sexangulare Brid., heath tundra, subalpine; sterile; 790,87-949,88-11-3,88-15-9,98790. (2) Cir­ cummontane. P strictum Brid., hummocks in mires, near cliff bases, subalpine and mid-elevations; cfr.; 87-250, 87- 959, 98821,99066. (3) Circumboreal. Pseudoleskea baileyi Best & Grout, heath tundra, subalpine; sterile; 98697, 98808, 87-789, 87-989, 88-6-27,88-6-27,88-7-31 ,88-8-35,88-9-39, 88-10-33, 88-66-33, 88-70-26. (1) Montane-West­ ern North America Endemic. P incurvata (Hedw.) Loeske , conglomerate outcrop, mineral soil of open slope, subalpine; sterile; 98685, 98744, 88067-X-l. (4) Circumboreal. P patens (Lindb.) Kindb., on cliff bases, lower elevations; sterile; 98686, 98979, 99036, 87-40-13, 87-3-7. (2) Montane-Wide Disjunct. P stenophylla Ren. & Card., epiphytic on reclining trunks of alder and on cottonwood, rock faces, at all elevations; cfr. ; 98719, 98751, 98966, 87-11-5, 87-11-24, 87-109, 87-796, 88-24-10, 88- 1310B, 87-12 7, 87-37-13, 1381B. (I) Temperate-North America Endemic Disjunct. Pseudotaxiphyllum elegans (Brid.) Iwats., cliff bases, mineral soil of slopes, lower elevations; sterile; 99136,87-151,87-191 , 99136. (2) Circumtemperate. Pterigynandrum filiforme Hedw., trunk of cottonwood, lower elevations; sterile; 98841. (4) Circum­ boreal. Ptilium crista-castrensis (Hedw.) De Not., open Sitka spruce stand, lower elevations; sterile; 87-20- W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 109

10. (4) Circumboreal. Pylaisiella polyantha (Hedw.) Grout, trunks of cottonwood, lower elevations; cfr.; 821, 1381C, 1381N, 98844, 99010, 99017. (4) Circumboreal. P. selwynii (Kindb.) Crum, Steere & Anderson, trunks of cottonwood, lower elevations; cfr.; 98937, 98996. (3) Circumtemperate. Racomitrium aciculare (Hedw.) Brid., boulders in streamlets, lower elevations, cfr.; 99173. (2) Cir­ cumboreal. R. affine (Schleich. ex Web. & Mohr) Lindb., slope edge of snowbed, boulders of boulder slope, all el­ evations; sterile; 99083, 99094, 99098, 99212, 730. (3) Temperate-Wide. R. canescens (Hedw.) Brid., snowbed margin, coastal cliffs and boulders, all elevations; sterile; 98688, 98689, 98707,98798, 98827,98894, 98668, 99898, 99193, 87-413, 87-420, 87-908. (3) Circumtemperate. R. fasciculare (Hedw.) Brid., boulders and outcrops in tundra, snowbed slopes, boulder slopes, all el­ evations; sterile; 98765, 98773, 98816, 99203, 87-770, 88-14-10. (3) Circumboreal. R. heterostichum (Hedw.) Brid., coastal boulder, lower elevations; sterile; 98094, 98098, 99083, 87- 417. (4) Circumboreal. R. macounii Kindb. , tundra, subalpine; sterile; 87-998. (3) Circummontane. R. muticum (Kindb.) Frisv., mineral soil, tundra slopes, subalpine; sterile; 98770, 98724, 99100. (3) Montane-AmphiPacific . R. occidentale (Ren. & Card.) Ren. & Card., base of boulder in lichen tundra, subalpine; sterile; 87- 792. (4) Temperate-Western North America Endemic. R. sudeticum (Funck) Brunch & Schimp., boulders, over rocks in tundra and near snowbeds, sub­ alpine; cfr.; 98748, 87-1022, 87-52, 87-856A, 87-859, 87-866, 87-869A, 87-927, 87-930A, 88- 10-30, 88-11-4, 88-11-9, 88-12-9, 88-13-10, 88-14-11, 88-14-12, 88-15-13, 88-15-17, 88- 1278B, 88-1500A, 87-997, 87-998, 98-754, 98-788. (1) Circumboreal. Rhizomnium glabrescens (Kindb.) T. Kop., humus banks, subalpine; sterile; 99177, 87-258. (3) Tem­ perate-Western North America Endemic. R. magnifolium (Horik.) T. Kop., snow-melt meadow, along brook in alder thicket, all elevations; ster­ ile; 98810, 98964. (4) Circumboreal. R. nudum (Britt. & Williams) T. Kop., alder thickets, heath tundra, Sitka spruce stand, all elevations; sterile; 98756, 98836, 98985, 87-172, 87-190, 87-190, 87-20-2, 87-20-26, 87-885, 87-975, 88- 46-4, 87-25-5. (3) Montane-AmphiPacific. Rhytidiadelphus loreus (Hedw.) Warnst., Sitka spruce forest and stream canyons, lower elevations; sterile; 99150, 87-458. (4) Boreal-Interrupted. R. squarrosus (Hedw.) Warnst., tundra and streambed margins, all elevations; sterile; 98781, 98890, 99091 , T3219-12, 87-873, 87-878, 88-1209A , 87-2-19, 87-19-15, 87-20-22. (1) Boreal-Inter­ rupted. R. triquetrus (Hedw.) Warnst., shady stream canyon, heath tundra, all elevations; sterile; 99099, 87- 511,87-521. (4) Circumboreal. Rhytidiopsis robusta (Hedw.) Broth., herb and heath tundra, subalpine; sterile; 98809, 87-971, 88-6- 29,88-7-29,88-8-27,88-67-25. (4) Montane-Western North America Endemic. Rhytidium rugosum (Hedw.) Kindb., escarpment top, subalpine; sterile; 98799. (4) Arctic-Alpine. Sanionia orthothecioides (Lindb.) Loeske, grassy cliff ledge, near sea level; sterile; 98863. (4) Arc­ tic-Circurnpolar. S. uncinata (Hedw.) Loeske, epiphytic on trees and shrubs, cliffs and bases, at all elevations; sterile; 98965, 98936, 98863, 99128, 87-108, 87-115, 87-126, 87-20-40, 87-21-16A, 88-1375N, 87-786, 110 J. Hattori Bot. Lab. No. 92 2 002

87-795, 88-1381A, 87-14-1, 87-14-2,87-14-4,88018-21, 87-78,87-410,87-415, 87-428, 87- 513. (I) Cosmopolitan. Sarmenthypnum sarmentosum (Wahlenb.) Tuom. & T. Kop. , depression in fen, mid-elevations; ster­ ile; 99084, 88-45-16, 88-45-17. (4) Arctic-Alpine. Schistidium boreale Poel!, cliffs and bolders, lower elevations, cfr.; 98667, 98747, 98867, 98870, 98891,99018,99141 . (3) Boreal-North America/Europe Disjunct. S dupretii (Ther.) W. A. Weber, conglomerate outcrop near snowbed, subalpine; cfr.; 98694. (4) Cir­ cummontane. S maritimum (Turn.) Bruch & Schimp., maritime outcrops, near sea level; cfr.; 98866, 98924, 99004. (4) Boreal-Interrupted. S. papillosum Culm in Amann, conglomerate cliff, near sea level; cfr.; 98667. (4) Circummontane. S platyphyllum (Mitt.) H. Perss., boulder in lichen tundra, subalpine; cfr.; 766 (4) Arctic-Alpine. S rivulare (Brid.) Podp., boulder in waterfall spray zone, rocks of stream margin, lower elevations; cfr.; 98867, 98870, 98891,87-585,43,37. (2) Boreal-Interrupted. Sphagnum angustifolium (C. Jens.) C. Jens., mires, mid-elevation; sterile; 88-60-22, 87-24-12, 87- 244, 87-250,87-23-10, 88-59-X-4. (I) Circumboreal. S annulatum H. Lindb., Sphagnum sedge mire, mid-elevation; sterile; 88-61-17. (4) Circumboreal. S balticum (Russ.) C. Jens., pool margin and in grass or sedge mires, mid-elevations; sterile; 88-48- X-3, 88-58-X-1, 88-59-X-3, 88-62-5. (3) Circumboreal. S compactum DC., seepage and in heath tundra, also in mires, subalpine and in mid-elevations; ster­ ile; 98789, 88-9-24, 88-9-25, 88-67-27, 87-345 87-347, 87-349, 87-995, 87-1008. (3) Circum­ boreal. S fimbriatum Wils., streamlet margin and grassy or sedge mires, mid-elevations; sterile; 88-48-X-6, 88-60-19, 88-59-17. (3) Circumboreal. Sfuscum (Schimp.) Klinggr. , mires, mid-elevations; sterile; 87-337, 87-353. (3) Circumboreal. S girgensohnii Russ., tundra, mires, streamlet banks, swamps, all elevations; sterile; 87-451, 87-452, 87-454, 87-513A, 87-516, 87-983, 87-984, 87-994, 87-996, 87-23-4, 87-25-4, 88-1295D, 88-32- 14,88-32-19. (\) Circumboreal. S. lindbergii Schimp., sedge mires, mid-elevations; sterile; 87-329, 87-330B, 87-343, 88-60-14. (3) Circumboreal. S. obtusum Warns!. , poor margin in mire, mid-elevation; sterile; 87-340. (4) Circumboreal. S papillosum Lindb., sedge-Sphagnum mires, mid-elevations; sterile; 88-61-15, 87-328, 87-330A, 87-334,87-335,87-336,87-348,87-331,87-344, 88-6/-16, T3230-11 . (4) Circumboreal. S riparium Angstr., sedge-Sphagnum mires, mid-elevations; sterile; 88-1312,88-7-15, 88-48-X-4, 88-58-12, 88-58-13, 88-58-X-4, 88-58-X-5, 88-60-13, 88-59-13, 88-60-16, 88-60-18, 88-60-21, 88-61 -X-2, 88-61 -X-3, 87-267. (I) Boreal-Interrupted. S russowii Warns!., sedge-Sphagnum mires, subalpine and mid-elevations; sterile; 88-48-X-7, 88-59- 14, 88-59-14A , 88-60-12, 88-60-20, 88-62-12, 88-62-13, 87-327, 87-332, 87-23-8, 87-337, 87- 338,87-341,87-350,87-352,87-999. (I) Boreal-Interrupted. S squarrosum Crome, sedge-Sphagnum mires, mid-elevations; sterile; 88-58-X-2, 88-58-X-3, 88-59- 15, 88-1295K, 87-269. (I) Circumboreal. S. subsecundum Nees, sedge-Sphagnum mires, mid-elevations; sterile; 99807, 87-264, 87-266A , 87- 268,87-2 70,87-22-51 . (3) Cosmopolitan. S teres (Schimp.) Angstr. , mires, mid-elevations; sterile; 87-257, 87-266, 87-268A, 88-48-X-2. (2) Circumboreal. S warnstorfii Russ. , heathland and mires, subalpine and mid-elevations; sterile; 98725, 98726, W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska III

Map 9. North American distribution of Tetrodontium repandum (Funck.) Schwaegr.

98771, 98779, 88-45-10, 88-47-11, 88-48-16, 88-48-24, 87-248, 87-265, 87-24-13, 87-24-18. ( I) Circumboreal. Tetraphis pellucida Hedw., rotten stump, Sitka spruce stand, low elevation; efr.; 87-469, 98988. (4) Circumboreal. Tetraplodon mnioides (Hedw.) Bruch & Schimp., intermittent streambed, subalpine; efr.; 87-881. (4) Circumboreal. Tetrodontium repandum (Funck) Schwaegr., shaded conglomerate cliff, sub alpine; efr.; 98763. (4) Boreal-Interrupted. (Map 9) Timmia austriaea Hedw., humid cliff crevices, sUbalpine and at sea level; sterile; 98714, 98768, 98901. (4) Arctic-Alpine. r norvegiea Zett., seepy cliff, near sea level; sterile; 98905. (4) Arctic-Alpine. Tomenthypnum nitens (Hedw.) Loeske, mires, mid-elevations; sterile; 99110, 87-246, 87-249, 88-48- 22. (4) Circumboreal. Tortellafragilis (Hook. & Wils.) Limpr., shaded dry maritime cliff, near sea level; sterile; 99146. (4) Circumboreal. r tortuosa (Hedw.) Limpr., cliff faces, lower elevations; sterile; 99052, 99179, 87-518. (4) Circum­ boreal. Tortula deciduidentata (Sharp & lwats.) Zander, humid sea cliff crevice, sea level; efr.; 98889. (4) Temperate-Western North America Endemic (Alaska and King Christian Island, Canada). (Map 10) r mueronifolia Schwaegr., maritime cliff, near sea level; efr.; 99023. (4) Circumboreal. r norvegiea (Web.) Wahlenb., forb meadow, mid-elevations; sterile; 88-38-18. (4) Arctic-Alpine. 112 1. Hattori Bot. Lab. No. 92 2 002

Map 10 (left). World distribution of Tortula deciduidentata (Sharp & Iwats.) Zander, a western North American endemic. Map II (right). World distribution of Schofieldia monticola Godfrey, a western North American endemic (excluding the Arctic locality).

T ruralis (Hedw.) Gaertn. et aI., cliff shelves, near sea level; sterile; 98869, 87-694, 87-702, 87-67, 87-134, 87-411 . (2) Circumboreal. Trematodon ambiguus (Hedw.) Hornsch., mineral soil at beaver pond, mid-elevations; cfr. ; 88-1300. (4) Boreal-Interrupted. Vlota drummondii (Hook. & Grev.) Brid., epiphytic on cottonwood and alder, all elevations; cf r.; 98932, 99005,98939, 88063-Y-J3. (3) Boreal-Interrupted. U obtusiuscula C. Miill. & Kindb., epiphytic on cottonwood and alder, all elevations; cfr. ; 87-40-39, 87-13-2,87-129, 87-130, 98879, 99003 . (3) Temperate-Western North America Endemic. U phyllantha Brid., dwarf shrub tundra, subalpine; sterile; 87-724. (4) Boreal-Interrupted. Warn storfia exannulata (Schimp.) Loeske, submerged in mire lagg in sedge fen , mid-elevations; ster­ ile; 99069,99080, 99081, 991 26, 99127, 99130, 87-360. (3) Circumboreal. W fluitans (Hedw.) Loeske, alder thicket depression, mid-elevation; sterile; 87-807. (3) Circumbore­ al. W pseudostraminea (Miill.) Tuom. & T. Kop. , willow thicket, mid-elevation; sterile; 88-45-X-1 . (4) Boreal-Interrupted. W trichophylla (Warnst.) Tuom. & T. Kop. , stream pool in mire, mid-elevations; sterile; 280, 87002- 9. (3) Boreal-AmphiPacific. W tundrae (Ameli) Loeske, floating in mire pond, mid-elevations; sterile; 99092. (3) Circumboreal.

Liverworts (56 taxa) Antheliajulacea (L.) Dum., mineral soil, seepage by streamlet, subalpine; sterile; 98786. (2) Boreal­ Interrupted. A. juratzkana (Limpr.) Trev., on soil, late snow sites, subalpine; sterile; 98690, 98774, 98791A. (2) W. B. SCHOF1ELD ET AL.: 8ryophytes from Tuxedni Wilderness area, Alaska 113

Circummontane. Barbilophoziajioerkei (Web. & Mohr) Loeske, heath tundra, subalpine; sterile; 99101, 98698, 98824. (2) Circummontane. B. hatcheri (Evans) Loeske, heath tundra, subalpine; sterile; 87-732, 98700, 98701. (3) Circumboreal. B. lycopodioides (Wallr.) Loeske, heath tundra, subalpine; sterile; 98811, 98832, 99206, 99215, 88- 18-14,88-21-10,88-22-13. (3) Circumboreal. Blasia pusilla L., moist gravel, low elevation; sterile; 87-567. (4) Circumboreal. Blepharostroma trichophyllum (L.) Dwn., shaded moist sites, lower elevations; sterile; 87-554, 99027. (4) Circumboreal. Calypogeia mulleriana (Schiffn.) K. Miill., shaded humus banks, low elevations; sterile; 99178. (4) Circumboreal. Cephalozia ambigua C. Massal., peaty opening in fen, subalpine; sterile; 99060. (4) Arctic-Alpine. C. bicuspidata (L.) Dwn., rotten stump in Sitka spruce forest, low elevation; sterile; 99020. (4) Cir­ cumboreal. C. catenulata (Hueb.) Lindb., shaded stream sites, lower elevations; clr.; 99174, 99176. (4) Circwn­ boreal. C. lunulifolia (Dum.) Dum., rotten logs, lower elevations; clr.; 98990, 99182. (4) Circumboreal. Chiloscyphus polyanthos (L.) Corda, splashed boulder in stream, lower elevations; sterile, 99038. (4) Circumboreal. Cladopodiellajiuitans (Nees) Buch, depression in fen, subalpine; sterile; 99073. (4) Circumboreal. Conocephalum conicum (L.) Lindb. & Mohr, shaded wet sites, lower elevations; sterile; 99134, 98750,87-197,87-591,87-602. (4) Circumboreal. Diplophyllum albicans (L.) Dum., mesic stream canyon, low elevation; sterile; 87-548. (4) Boreal-In­ terrupted. D. plicatum Lindb., edge of snowbed, subalpine; sterile; 98733. (4) Boreal-Amphi Pacific. D. taxifolium (Wahlenb.) Dum., shaded cliffs and mineral soil, all elevations; sterile; 98678, 98728. (3) Circumboreal. Gymnomitrion concinnatum (Lighft.) Corda, cliff faces, subalpine; sterile; 98712, 98730, 98806, 99115. (3) Montane-Interrupted. G. obtusum (Lindb.) Pears., dry cliffs, subalpine; sterile; 98758. (4) Temperate-Wide. Jungermannia confertissima Nees, seepy cliff, low elevations; sterile; 87-597. (3) Circumboreal. J. exsertifolia Steph., damp edge of snowbed, on rock, subalpine and near sea level; sterile; 99113, 586, 594, 609. (3) Arctic-Alpine. J. obovata Nees, damp cliffs, lower elevation; sterile; 99162, 87-602. (3) Circwnboreal. Lophocolea heterophylla (Schrad.) Dum., rotten logs, lower elevations; cfr.; 98950, 98951, 87048-7. (4) Circumboreal. Lophozia badensis (Gott. ex Rabenh.) Schiffn., mineral soil, stream canyon, lower elevations; sterile; 517. (4) Circumboreal. L. bicrenata (Schrnid.) Dum., heath tundra, higher elevations; cfr.; 99103 . (4) Circwnboreal. L. incisa (Schrad.) Dum., rotten log and humus, lower elevations; sterile; 98993, 99170. (3) Circum- boreal. L. opacifolia Culm., shaded cliff; sterile; 98680. (4) Montane-Interrupted. L. ventricosa (Dicks.) Dum., rotten logs, lower elevations; sterile; 98991,231. (4) Circumboreal. L. wenzelii (Nees) Steph., cliff top, subalpine; sterile; 98705. (4) Arctic-Alpine. Marsupella alpina (Gott.) Bemet, heath tundra, terrestrial, subalpine; sterile; 98730B, 99104. (4) Arctic-Alpine. 114 1. Hattori Bot. Lab. No. 92 2 0 0 2

M. brevissima (Dum.) Grolle, snowbed slopes, subalpine; sterile; 98691, 88011-11, 88014-9, 88-11- 5,88-13-8. (4) Arctic-Alpine. M. emarginata (Ehrh.) Dum., moist rock surfaces, all elevations; sterile; 98676, 87-516, 87-718A, 88-1295A . (3) Circumboreal. M. sparsifolia (Lindb.) Dum., snowbed edge, subalpine; sterile; 99122. (3) Montane-Interrupted. M. sphacelata (Gies.) Dum., snowbed edge, subalpine; sterile; 99123, 99124, 88-1295A . (3) Mon­ tane-Interrupted. Moerckia blyttii (Moerck) Brockm. streamlet margin, subalpine; sterile; 98683, 98769, 87-273. (3) Montane-Interrupted. Mylia anomala (Hook.) S. Gray, in Sphagnum hummocks in fen, mid-elevation; sterile (gemmae); 99070. (4) Circumboreal. Nardia scalaris S. Gray, streamlet bed, lower elevation; sterile; 1295M. (4) Circumboreal. Pellia endiviifolia (Dicks.) Dum., on rock, lower elevations; sterile; 87-696. (4) Circumboreal. P neesiana (Gott.) Limpr., shaded banks and cliff bases, all elevations; sterile; 98679, 98767, 99155, 917. (4) Circumboreal. Plagiochila porelloides (Torr.) Lindb., shaded cliffs, lower elevations; sterile; 98860, 99048. (4) Cir­ cumboreal. Pleurocladula albescens (Hook.) Grolle, snowbed edge, subalpine; sterile; 98729, 98730A, 87-880. (3) Arctic-Alpine. Porella cordaeana (Hueb.) Moore, rock faces, shaded, lower elevations; sterile; 98850, 98853, 98861, 99053, 87-706. (3) Boreal-Interrupted. Preissia quadrata (Scop.) Nees., shaded humid cliffs, lower elevations; cfr.; 98904, 99200. (4) Cir­ cumboreal. Ptilidium ciliare (L.) Hampe, shrub tundra, subalpine; sterile; 87-722, 87-820A, 88-70-28, 88-73-20, 99203A. (3) Circumboreal. P pulcherrimum (G. Web.) Vainio, epiphytic on Sitka spruce trunks, lower elevations; sterile; 98982, 98989,87-232,88-137511, 88-1376F, 88-13761. (3) Circumboreal. Radula complanata (L.) Dum., shaded cliff, lower elevations; cfr.; 99051 , 87-14. (4) Circumboreal. Scapania americana K. Mull., cliff near waterfall, lower elevation; cfr.; 88A. (4) Temperate-Western North America Endemic. S. irrigua (Nees) Nees, hummock, edge offen, mid-elevation; sterile; 99107. (4) Circumboreal. S. subalpina (Nees.) Dum., in alder thicket and at edge of snowbed, lower and subalpine elevations; sterile; 99112,87-800. (4) Circumboreal. S. uliginosa (Sw. ex Lindenb.) Dum., intermittent streamlet, subalpine; sterile; 98677, 874. (4) Cir­ cumboreal. S. umbrosa (Schrad.) Dum., rotten log in cottonwood thicket, low elevation; sterile; 98938. (4) Bore­ al-Interrupted. S. undulata (L.) Dum., snowmelt streamlet, spray zone at waterfall, subalpine; 98780, 98787, 98677, 99022, 87-601A . (3) Circumboreal. Schofieldia monticola Godfrey, snowbed margin, subalpine; sterile; 98738, 98830, 98838. (3) Mon­ tane-Western North America Endemic. (Map 11) Tetralophozia setiformis (Ehrh.) Urmi, top of outcrop knob ledge, subalpine; sterile; 98822. (4) Cir­ cumboreal. Tritomaria quinquedentata (Huds.) Buch, on boulder, stream canyon, lower elevations; 98764A, 56. (4) Boreal-Interrupted. W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska liS

Remarks on Particularly Significant Species Antitrichia californica Sull. (Map 1) This species is one of a group of bryophytes that are disjunct between the Mediter­ ranean and Pacific North America. This collection appears to represent its northern limit in the Western Hemisphere, although it extends westward to Kodiak Island (Peterson et al. 1980) and Ivanof Bay (Schofield, unpublished data, 1997) on the Alaska Peninsula. Brachydontium olympicum (Britt.) McIntosh & Spence (Map 2) This species, originally described from Washington State, is infrequent northward in British Columbia and Alaska and southward to Mount Hood, Washington State. It occurs mainly on volcanic rock in alpine and subalpine late-snow areas, but on Chisik Island was on conglomerates. It is also rare in Japan. Bryum miniatum Lesq. (Map 3) This appears to be the first report from Alaska. Its presence in the Queen Charlotte Is­ lands of British Columbia strongly implies that the species would be expected in the Alexander Archipelago of Southeast Alaska. It was rare on Chisik Island. Calliergon megalophyllum Mik. (Map 4) This species, previously considered rare in North America, is shown to be widely dis­ tributed along the Alaska Peninsula and Aleutian Islands (Schofield, unpublished data, 1995- 2000). Dichodontium olympicum. Ren. & Card. (Map 5) This collection is the only one from Alaska, and represents its northernmost locality. The species is endemic to western North America, from California northward to Alaska. It is predominantly subalpine and grows on soil in well-drained sites. Ditrichum zonatum (Brid.) Lindb. This species is readily overlooked because it lacks sporophytes and closely resembles D. heteromallum. It is predominantly a subalpine species, and is known also from Europe. Echinophyllum sachalinensis (Lindb.) O'Brien in O'Brien & Horton (Map 6) This species superficially resembles Claopodium bolanderi Best, a species not infre­ quent in Southeast Alaska and southward along the Pacific Coast. In Alaska, Echinophyl­ lum appears to be confined to tundra. It was reported first from non-peninsular Alaska and has been found at a single locality in the Rocky Mountains of British Columbia (O'Brien & Horton 2000). Specimens lacked sporophytes. The species is disjunctive in East Asia, from where it was described. Hymenostylium insigne (Dix.) Podp. (Map 7) Plants of this species are always without sporophytes. It is confined to moist calcare­ ous cliffs, and appears to be more widespread in western North America than published re­ ports note. Collections have been seen from various sites in coastal British Columbia and Southeast Alaska. Hypnum cupressiforme Hedw. var. subjulaceum Mol. In the Tuxedni Wilderness Area, this moss was represented only by the variety. It ap­ pears to be relatively frequent on the Alaska Peninsula and associated islands as well as the Aleutian Islands. Iwatsukiella leucotricha (Mitt.) Buck & Crum 116 1. Hattori Bot. Lab. No. 92 2 002

Steere (1978) mapped the distribution of this species in Alaska, where he notes a wider ecological range than we have observed in Japan or southeastern Alaska and south­ ward to Oregon. There, it is mainly on conifers Abies, Chamaecyparis, and Pinus. In areas of the Alaska Peninsula (Schofield, unpublished data, 1995-2000), we have seen it only on Alnus and Populus and the most luxuriant populations are on Chisik Island, where it pro­ duced abundant sporophytes, the first that appear to have been noted in the North America material. Along much of the Pacific Coast of North America, the species is most frequent near sea level, but in Oregon it appears at higher elevations where it is rare. Mielichhoferia macrocarpa (Hook.) Bruch & Schimp. Brassard (1971) mapped the North American distribution of this species, showing a single locality in Alaska. It is also known from two localities in Southeast Alaska as well as several populations on Chisik Island. Oedipodium griffithianum (Dicks.) Schwaegr. Although relatively frequent in the Alexander Archipelago, the species is rare in the rest of Alaska, extending westward to Chisik and Kodiak Islands (Peterson et al. 1980). It is most frequent near sea level. Orthotrichum pallens Bruch This species is new to Alaska, and represents its northernmost locality in North Amer- ica. Rhytidiopsis robusta (Hedw.) Broth. This western North American endemic is widespread in alpine and sub alpine habitats in Oregon, northward to Southeast Alaska. It rarely occurs outward along the Alaska Peninsula (Schofield, unpublished data, 1995-2000); populations on Chisik Island and Mother Goose Lake on Alaska Peninsula are the northernmost and westernmost. It can be expected on the Kenai Peninsula and the Chugach Mountains. Schofieldia monticola Godfrey (Map 8) This species appears to be rare in Alaska. Schuster (1995) reported it from Girdwood area in the Chugach Mountains. The Chisik Island population extends it westward. It is likely to be present in the mountainous areas of Kodiak Island as well as on the Kenai Peninsula. Tetrodontium repandum (Funck.) Schwaegr. (Map 9) This species is probably rare throughout its interrupted range, although it is readily overlooked. It is likely to extend along the Aleutian Chain. Its presence here on conglomer­ ate is an unusual substrate. Tortula deciduidentata (Sharp & Iwats.) Zander (Map 10) This species, originally described from Port Moller on the Alaska Peninsula, appears to be widespread on that peninsula and associated islands (Schofield, unpublished data, 1995-2000). It was extremely rare on Chisik Island. Map 10 shows the Alaskan distribu­ tion; a single, isolated locality is known from King Christian Island in the Canadian arctic archipelago (Vitt and Zander 1978).

Bryophytes ofParticular Habitats Alder thickets (Fig. 3) provide shade for common terrestrial boreal species of W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 117

bryophytes and also for epiphytes. On branches, Pseudoleskea stenophylla, Ulota drum­ mondii, U obtusiuscula, U phyllantha, and a few other bryophytes are present, but never abundant. On the trunks, either erect or reclining, Brachythecium refiexum, Dicranum tau­ ricum, Iwatsukiella leucotricha, and a few other bryophytes are present, reaching their greatest luxuriance near sea level. At upper elevations, there is often a mosaic of open mineral soil and heaths. These sites harbored the following bryophytes: Andreaea alpestris, Barbilophozia fioerkei, Conostomum tetragonum, Desmatodon latifolius, Echinophyllum sachalinensis, Hypnum plicatulum, Kiaeria falcata, K. blyttii, Lophozia opacifolia, Marsupella brevissima, Olig­ otrichum hercynicum, Pseudoleskea bai/eyi, Racomitrium muticum, and R. fasciculare. Where snowbeds persisted in the tundra or over outcrops irrigated by snow-melt streams, the characteristic alpine-subalpine bryophytes were present, including Andreaea blyttii, A. nivalis, Anthelia juratzkana, Arctoa fulvella, Brachydontium olympicum, Bryum weigelii, Diplophyllum plicatum, Ditrichum zonatum, Gymnomitrion concinnatum, Hyp­ num callichroum, Marsupella sphacelata, M. sparsifolia, Moerckia blyttii, Pleurocladula albescens, Pseudoleskea baileyi, Oligotrichum hercynicum, 0. parallelum, Racomitrium fasciculare, R. muticum, R. sudeticum, Rhytidiopsis robusta, Rhizomnium nudum, and Schofieldia monticola. Most of the mires were provided by at least some ground-water, and in some it was undoubtedly the main source. In consequence, there is a mixture of bog and fen species. All peatlands were sha\low with few sma\l pools. Sphagnum riparium appears to be the most frequent species, but other species including S. balticum, S. papil/osum, S. russowii, S. subsecundum, S. teres, and S. warnstorfii, etc. are also present. Other mosses in these wetlands include Calliergon stramineum, C. sarmentosum, Campylium stellatum, Cinclidi­ um subrotundum, Dicranum undulatum, Hypnum lindbergii, Polytrichum strictum, Pohlia sphagnicola, and Warnstorfia exannulata. In one wetland Paludella squarrosa was present. Populus trichocarpa stands, occasional at lower elevations, have their own suite of bryophytes on the stone-like bark: Antitrichia curtipendula, Brachythecium reflexum, Pterigynandrum filiforme, Pseudoleskeella nervosa, Pylaisiella polyantha, and several species of Orthotrichum, including 0. obtusilobum. Decaying logs also provide sites for bryophytes essentially restricted to such sites and infrequent on the island: Cephalozia bicuspidata, C. lunulifolia, Lophocolea heterophylla, Lophozia incisa, Mnium spinulosum, Scapania umbrosa, Tetraphis pellucida, etc. Sitka spruce stands, extremely local on the islands, and very limited in extent, harbored the epi­ phytes Isothecium stoloniferum, Oncophorus wahlenbergii, and Ptilidium pulcherrimum, not found elsewhere. The floor of this relatively open "forest" had a few bryophytes not noted elsewhere, including Dicranum majus, Plagiothecium undulatum, and Plagiomnium insigne. Although nesting seabirds greatly restrict variety of bryophytes on the cliffs they in­ habit, other cliffs showed some diversity. Near sea level drier cliffs possessed Antitrichia californica, Encalypta ciliata, E. rhaptocarpa, Herzogiella adscendens, Hypnum cupressi­ forme var. subjulaceum, Porella cordaeana, etc. Drier cliffs at higher elevations had Abi­ etinella abietina, Andreaea rupestris, Arctoa fulvella, Rhytidium rugosum, Tetralophozia 118 1. Hattori Bot. Lab. No. 92 200 2 setiformis, etc. Wet, usually somewhat shaded cliffs produced greater luxuriance and diversity of bryophytes. These cliffs were near sea-level and continuously irrigated by seepage or formed the mouth of streamlets. Here were Amphidium lapponicum, Anomobryum filiforme, Blindia acuta, Brachythecium rivulare, Bryoerythrophyllum inaequalifolium, Cir­ riphyllum cirrosum, Cratoneuron filicinum, Distichium capillaceum, Gymnostomum aerug­ inosum, Hygrohypnum bestii, Hymenostylium insigne, H. recurvirostre, Myurella julacea, Philonotis Jontana, Preissia quadrata, Timmia austriaca, and T norvegica. Oedipodium griffithianum occurred locally on shaded cliffs both near sea level as well as at higher ele­ vations. Small streams dissect the island, most of them having their water source from persis­ tent snow-beds of higher elevations. These have excavated shallow canyons. Their flanking rock outcrops provided sites for bryophytes infrequent elsewhere on the island, and never common in the canyons: Bartramia ithyphylla, B. pomiformis, Bartramiopsis lescurii, Fis­ sidens bryoides, Herzogiella striatella, Isopterygiopsis pulchella, Radula complanata, Rhi­ zomnium glabrescens, etc. On the boulders or streambed rock in the streams occurred Hy­ grohypnum alpinum, H. ochraceum, and Schistidium rivulare. Human influence on the bryophytes was apparent only near the old cannery where Leptobryum pyriforme and Pohlia nutans were abundant. Of particular interest was Funar­ ia hygrometrica that was confined to shaded cliff strata near the sea, and not influenced by human activity. Marchantia polymorpha, too, was clearly not introduced by human activity.

Analysis of the Flora An analysis of the mosses into geographic categories revealed 61 % are boreal species, which clearly dominate the flora (Table 1). The second largest category, montane, account­ ed for 13% of the flora. The distinction between boreal and temperate reflects the generally more southern distribution of most species; montane species are subalpine on the island; endemic North American species are predominantly those of the Pacific Coast, although a few extend across North America. The preponderance of boreal species that are widespread in the northern hemisphere is a response to the relatively mild temperatures found on the island. The combined tem­ perate and boreal elements is 72%. Arctic-Alpine and montane species reflect the relatively limited area of suitable habitats available; much of the island is vegetated by a tangle of alder thickets that extend from the lower limits of the sub alpine to near sea level. Peatland is limited and scattered at mid-elevations. Endemic western North American species, 4% of the total flora, are found at all elevations; a third are montane through their total range, but are subalpine on Chisik Island. Frequency of occurrence, using the four categories defined in the "Checklist Format" section, is shown in Table 2, indicating that a very high proportion of the species are either uncommon or rare on the island (82%) and the residue (18%) common or abundant. The relative uniformity of much of the environment is conspicuous and low diversity is a conse­ quence, while the many micro-habitats of the subalpine, stream canyons, and sea level habitats harbor the greatest diversity ofbryophytes. W. B. SCHOFIELD ET AL.: 8ryophytes from Tuxedni Wilderness area, Alaska 119

Table 1. Percent distribution of Tuxedni Wilderness Area, Alaska, bryophyte species in phytogeographic categories.

Phytogeographic category Number of species Percent of flora

ARCTIC 21 7.3 Arctic-Alpine 19 6.6 Arctic-Circumpolar 2 0.7 BOREAL 173 60.5 Circumpolar 131 45 .8 Borea1-Interrupted 33 11.5 Boreal-AmphiPacific 7 2.8 Boreal-North America/Europe Disjunct 1 0.4 TEMPERATE 34 11.9 Temperate-Western North America Endemic 8 2.8 Circumtemperate 8 2.8 Temperate-Interrupted 7 2.5 Temperate-Western North AmericalEurope Disjunct 6 2.1 Temperate-Wide 3 1.1 Temperate-AmphiPacific I 0.4 MONTANE 38 13.3 Circummontane 20 7.0 Montane-Interrupted 5 1.8 Montane-Western North America Endemic 5 1.7 Montane-Wide Disjunct 4 1.4 Montane-AmphiPacific 4 1.4 COSMOPOLITAN 20 7.0 Total 286 100.0

Table 2. Frequency of occurrence of moss species in four abundance categories within Tuxedni Wilderness Area, Alaska.

Commonness category Number of species Percent of flora

Rare 146 51.1 Uncommon 87 30.4 Common 33 11.5 Abundant 20 7.0 Total 286 100.0

To suggest an answer to the possibility that reproduction of readily dispensable dias­ pores might be the main control of the frequency of taxa, we compared the number of species with sporophytes with those that lack them (Table 3). Species commonly with sporophytes account for 34% of the flora, while those without them are 66%. Looking at 120 1. Hattori Bot. Lab. No. 92 2 0 0 2

Table 3. Comparison of the numbers of bryophytes from Tuxedni Wilderness Area, Alaska, always lacking sporophytes with those that are commonly with sporophytes.

Category Number of species Percent of flora

Always lacking sporophytes 187 65.4 Commonly with sporophytes 99 34.6 Total 286 100.0

specific cases, many taxa that lacked sporophytes were rather widespread on the island, while many with sporophytes were rather local. It seems likely, however, that bryophytes that reached the island from the mainland may be equally probable to have arrived as vege­ tative propagules as through spores. Frequent re-introduction seems probable since the is­ land is close to the mainland. It seems probable that coincidence of reaching suitable sites to colonize would be the predominant factor in their establishment, and the limitation of suitable, and already colonized, sites would hamper more recent colonizations in spite of possible arrival of diaspores. Although most bryophytes on Chisik Island are widespread circumpolar species, sev­ eral represent species that show a highly disjunctive pattern in the Northern Hemisphere: Andreaea nivalis, Anoectangium aestivum, Antitrichia californica, A. curtipendula, Bry­ oerythrophyllum inaequalifolium, Bryum miniatum, *Cynodontium jenneri, *Dicranum tauricum , *Ditrichum zonatum, *H ymenostylium insigne, Oedipodium griffithianum, Tetrodontium repandum, and Vlota drummondii among the mosses, and *Lophozia opaci­ folia , Marsupella alpina, and Porella cordaeana among the hepatics. Those marked with an asterisk (*) are relatively widespread, and sometimes common in Europe. North Ameri­ can endemics include the following: Brachythecium hoizingeri, Claopodium bolanderi, Di­ chodontium olympicum, Hygrohypnum bestii, Isothecium stoloniferum, Mielichhoferia macrocarpa, Plagiomnium insigne, Pseudoleskea baileyi, Racomitrium occidentale, Rhi­ zomnium glabrescens, Rhytidiopsis robusta, Schofieldia monticola, and Tortula deciduiden­ tata. Those that follow an arc around the North Pacific, mainly near the coast, to Southeast Asia, include the mosses Bartramiopsis lescurii, Brachydontium olympicum, Bryhnia hul­ tenii, Echinophyllum sachalinensis , Herzogiella adscendens, Iwatsukiella leucotrichia, Oligotrichum aligerum, Oligotrichum parallelum, Pseudoleskea stenophylla, Racomitrium muticum , and Rhizomnium nudum, and the hepatic Diplophyllum plicatum. Each of these patterns represents a fragment of a richer assemblage in the most ocean­ ic portions of Southeast Alaska and coastal British Columbia. All of these taxa can be speculated to have extended northward postglacially from periglacial refugia into Alaska and now persist in favorable sites that have not been invaded by vascular plants or by more aggressive bryophyte taxa. Most were probably survivors in unglaciated North America, but others, for example Bryhnia huitenii, may have extended their range from Southeast Asia along the Aleutian Chain and the Alaska Peninsula, where they are relatively wide- W. B. SCHOFIELD ET AL.: Bryophytes from Tuxedni Wilderness area, Alaska 121 spread. The western North American endemic Tortula deciduidentata shows a distribution confined to calcareous substrata near the ocean. This suggests that it may have survived glaciation within the area and then expanded its range. It is rather scarce through much of its restricted range, but produces sporophytes in abundance, so may be restricted more by the infrequency of a suitable habitat than by its potential dispersibility.

ACKNOWLEDGMENTS Financial and logistical support were provided by the Air Quality Branch, Division of Refuges, U.S. Fish and Wildlife Service, Denver, and the Alaska Maritime National Wildlife Refuge, Homer. We gratefully acknowledge the field assistance of Nils E. Talbot during the 1987 season, Richard E. Andrus for determining Sphagnum, Stanley L. Welsh for determining vascular plants, John G. Brewer for map design and production, and the staff of Columbia-Ward Fisheries for lodging and support, and Michael S. Ignatov and Norton G . Miller for thoughtful reviews of the manuscript.

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