The Bryologist 105(1), pp. 111 121
Copyright q 2002 by the American Bryological and Lichenological Society, Inc.
Lichens from Simeonof Wilderness, Shumagin Islands, Southwestern Alaska
STEPHEN S. TALBOT
U.S. Fish and Wildlife Service, 1011 East Tudor Road, Anchorage, AK 99503, U.S.A.
SANDRA LOOMAN TALBOT
Alaska Science Center, U.S. Geological Survey, Biological Resources Division, 1011 East Tudor Road,
Anchorage, AK 99503, U.S.A.
JOHN W. THOMSON
Department of Botany, University of Wisconsin, Madison, WI 53706, U.S.A.
FRED J. A. DANIËLS
Institute of Plant Ecology, Westfälische Wilhelms-Universität, Hindenburgplatz 55,48143 Münster, Germany
WILFRED B. SCHOFIELD
Department of Botany, University of British Columbia, Vancouver, British Columbia V6T 1Z4, Canada
Abstract. One hundred eighty-eight taxa of lichens are reported from Simeonof Island in the
Shumagin Islands of southwestern Alaska. Wide-ranging arctic-alpine and boreal species dominate
the lichens; a coastal element is moderately represented, while amphi-Beringian species form a
minor element. The lichen component of Empetrum nigrum dwarf shrub heath, the dominant
vegetation type, was analyzed to identify the most frequently occurring lichens within this community.
Simeonof Wilderness, located within the Shumagin Islands south of the Alaska Peninsula, Alaska, includes Simeonof Island and the Murie Islets
and lies within the Western Pacific Coast phytogeographic district of Hultén (1941–1950) (Fig. 1). The
Simeonof landscape is diverse with small mountains, oceanic heaths, meadows, deciduous thickets,
lakes, streams, poor fens, and sandy and cobbly
beaches (Fig. 2). As part of the Alaska Maritime
National Wildlife Refuge managed by the U.S. Fish
and Wildlife Service, Simeonof Wilderness is a
Class I air quality area (Clean Air Act, 42 U.S.
Code 7401 et seq.). It is also a National Natural
Landmark administered by the U.S. National Park
Service (Swain 1990). In accordance with its responsibility for ensuring high air quality standards
in Simeonof Wilderness, the Service sought to assess the existing condition of wilderness resources
of these islands relative to air quality using the
guidelines proposed by Fox et al. (l987), in which
the first step in characterizing the biological components is to document the lichens with habitat and
distributional information. As the lichen diversity
of Simeonof Wilderness—and of the islands south
of the Alaska Peninsula region—is virtually unknown (Krog 1968; Thomson 1984, 1997), the primary objective of the present study was to list the
lichens, including habitat and distribution information, for Simeonof Wilderness.
STUDY AREA
The Shumagin Islands are situated about 900 km southwest of Anchorage south of the Alaska Peninsula. From
east to west, the islands span 100 km and include Unga,
Popof, Korovin, Nagai, Chernabura, Big and Little Koniuji, and Simeonof Island plus other smaller islands. Simeonof Island is located in the southeastern corner of the
Shumagin Islands group at 548549 N, 1598189 W. Radiocarbon dating of peat from Unga and Popof Island suggests that deglaciation probably dates from 12,000 to
10,000 yr BP (Heusser 1983).
The Shumagin Islands are mountainous, with peaks of
the largest islands averaging 506 m elevation (698 m maximum on Unga Island). Unlike most of the mountains of
the Shumagin Islands whose steep slopes arise abruptly
from the sea, the mountains of Simeonof Island arise from
extensive lowlands dotted with numerous ponds and wetlands. The largest of the three mountains is 438 m high.
Bedrock of the northern and central Shumagin Islands
is of sandstone and conglomerate, some of which is volcanic and other plutonic rock, while the southern islands,
including Simeonof, are of granodiorite (Burk 1965). Between 12,000 to at least 9,000 B.P., the Shumagin Islands
formed a single, circular peninsula on the Shumagin Shelf
extending southeastward from the Alaska Peninsula. The
Shumagin Group became insular about 8,000 to 6,000 B.P.
(Winslow 1992).
The climate of the Shumagin Islands is maritime with
considerable wind and cool humid and cloudy conditions
(Heusser 1983). There are no available long-period climatological records for the Shumagin Islands. However,
the climatological records for a 42-yr period from Cold
Bay (558129 N, 1628429 W), approximately 180 km west
northwest of Simeonof, indicate a mean annual temperature of 3.38C and a mean annual precipitation is 871 mm
0007-2745/02/111–121$1.250/0
112
THE BRYOLOGIST
[VOL. 105
FIGURE 1. Location of Simeonof Island, Alaska, and the general geography of the island. Contour interval is 75
m. The elevation of the highest mountain is 508 m.
(Leslie 1989). Monthly maximum and minimum temperature variations average less than 128C and the moderating
effect of the ocean results in indistinct seasonal changes
(Searby 1959). Using the ecoclimatic-phytogeographical
system of Tuhkanen (1984), Cold Bay is in the northern
boreal subzone, hyperoceanic (O2) sector, and humid (h)
province. Climatic data for a shorter period (7 yr) are
available from Sand Point, Popof Island, Alaska (558209
N, 1608309 W), approximately 100 km northwest of Simeonof Island; these data show a warmer and wetter climate: mean annual temperature, 4.78C, and mean annual
precipitation, 1,199 mm (Leslie 1989). According to the
system of Tuhkanen (1984), Simeonof Island is located in
the middle boreal subzone, hyperoceanic (O2) sector, and
humid (h) province. This indicates that Simeonof Island
occurs in a warmer subzone than Cold Bay and accordingly has a longer growing season.
Soils of Simeonof Island are mainly Typic Cryandepts
(Rieger et al. 1979). These are well-drained, silt loam soils
with a dark-colored surface horizon ranging from seven
to 37 cm thick. Most of these soils have a smeary, loamtextured subsoil developed over stratified sandy and cindery volcanic deposits. They support both grass-alder and
shrubby tundra vegetation. Other soil types include Fluvaquentic Cryofibrists on very poorly drained organic
soils in depressions and broad valley bottoms, Lithic Cryofolists on high ridges, and Typic Cryopsamments on ex-
cessively drained soils on coastal dunes. If disturbed, the
latter are subject to wind erosion.
The vegetation of Simeonof Island is naturally treeless
(‘maritime tundra’) and primarily consists of natural coastal cliff-, meadow-, and dune-vegetation and maritime and
alpine dwarf shrub tundra vegetation. A variety of wetland
communities and forb-, willow- and alder vegetation grow
in sheltered sites. The zonal vegetation of Simeonof Island
is Empetrum nigrum dwarf shrub heath. An overview of
the vegetation is given in Daniëls et al. (1998).
METHODS
Lichen taxa. Field studies were conducted during September 1981, July 1995, and July 1996. A total of 781
specimens were collected. Most determinations were
made by the third author (JWT) and these are given first
in the list of lichens; some others were determined by
FJAD and these follow a semi-colon in the list. Voucher
specimens are deposited in the University of Wisconsin
Herbarium (WIS) and Westfälische Wilhelms-Universität
(MSUN). Nomenclature follows Esslinger and Egan
(1995) with Internet updates (Esslinger 1998).
Lichen collections were made in conjunction with other
botanical studies during the three summer sampling periods. In 1981, our primary duty was to assess the range
condition; in 1995, it was to monitor vegetation change
2002]
TALBOT ET AL.: SIMEONOF WILDERNESS LICHENS
113
FIGURE 2. View of the northern portion of Simeonof Island, Alaska, from the alpine zone of a mountain in the
center of the island. This image shows Simeonof Island composed of two separate main portions which remain connected by a sandy isthmus at the head of Simeonof Harbor. The elevation of the mountain on the north side of Simeonof
Harbor is 311 m. Other islands of the Shumagin Islands are seen in the background.
following the cattle removal; and in 1996, to collect lichen
and moss samples for elemental analysis in relation to air
quality and to describe and classify heath plant communities. During each of the three periods, collections of
macro- and microlichens were made to develop a baseline
list of taxa as we explored the island and traveled on foot
to randomly selected sample sites. In collecting lichens,
an attempt was made to include a representative spectrum
of habitat types over complex topographic gradients.
World distribution patterns are assigned to each taxon
using the following geographic categories (many have
combinations of these): A, arctic-alpine, north of the latitudinal treeline south into the mountains of the east (Adirondak and White Mountains) and west (Rocky Mountain
and Cascade Ranges) (Thomson 1984); B, boreal, latitudinal treeline to the southern limit of the closed coniferous
forest with some species also common in the temperate
regions; C, coastal, present in a strip along the maritime
coast and reflecting a maritime climate; M, amphi-Beringian, western American Arctic and also in eastern Asia;
W, widespread, present in many or most of the other categories; T, temperate; U, unclear distributions, either under study or too poorly known taxonomically to be confident of published maps. We assigned these phytogeographic categories based on Ahti (1980), Arup (1994),
Culberson and Culberson (1968, 1978), Dibben (1980),
Geiser et al. (1994), Gould (1994), Goward et al. (1994),
Jahns (1981), Søchting and Olech (1995), Talbot et al.
(1991, 1992, 1997), and Thomson (1984, 1997).
Lichen components of heath vegetation. Oceanic
crowberry (Empetrum nigrum) heaths are the dominant
plant community type of Simeonof Island (Daniëls et al.
1998). To identify lichens of high presence within crowberry heaths, we described the diversity of lichen composition along mesotopographic gradients from lowlands
to alpine. The study is based on a set of 30 relevés made
according to the Braun-Blanquet approach (Westoff & van
der Maarel 1973). Plots were laid out in units of homogeneous crowberry heath vegetation so as to represent
conspicuous environmental variation of the heath vegetation from lowlands to alpine. Relevé size, 25 m2, approximately equaled the minimal area for comparable types
(Westoff & van der Maarel 1973).
RESULTS
One hundred eighty-eight taxa were found in Simeonof Wilderness Area. We were unable to find
any references in the published literature regarding
the lichens of Simeonof. Thus, all 188 species are
new reports for the wilderness area. Of nine species
reported by previous authors from the Shumagin
Islands (Cummings 1910; Rothrock 1884; Thomson 1984), five, Micarea lignaria (Ach.) Hedl. (5
Biatora milliaria Fr.), Nephroma bellum (Sprengel)
Tuck., Pertusaria octomela (Norman) Erichsen,
Physcia stellaris (L.) Nyl., and Ramalina farinacea
(L.) Ach., were not collected by the present authors.
The remaining four species were collected by the
present authors: Mycoblastus alpinus (Schaerer)
Schauer (5 Heterothecium sanguinarium (Fl.)
Tuck. var. alpinum Fr.), Ochrolechia tartarea (L.)
Mass. (5 Lecanora tartarea (L.) Ach.), Peltigera
aphthosa (L.) Willd., and Sphaerophorus globosus
(Hudson) Vainio.
The distribution patterns for lichens from Simeonof Island are similar to those from Izembek
114
THE BRYOLOGIST
[VOL. 105
FIGURE 3. Geographic distribution patterns of lichens of Simeonof Island, Alaska in relation to other western Alaska
sites. Percent values indicate the presence of each taxon occurring within each category; some taxa are shared among
categories; n 5 total number of lichens (in parentheses) from Adak Island (Talbot et al. 1997), Attu Island (Talbot et
al. 1991), and Izembek National Wildlife Refuge (NWR) (Talbot et al. 2000).
National Wildlife Refuge in the westernmost portion of the Alaska Peninsula that extends to the
Aleutian Islands (Talbot et al. 2000), Adak Island
in the central Aleutian Islands (Talbot et al. 1997),
and Attu Island in the westernmost Aleutian Islands
(Talbot et al. 1991) (Fig. 3). The lichens of these
areas are characterized by wide-ranging, arctic-alpine and boreal species with a moderately well-represented coastal element. Amphi-Beringian species
form a minor element.
The name of each taxon is followed by habitat
information which is in turn followed by elevational range and collection number. Vascular plant taxonomy follows USDA, NRCS (2001). Collection
numbers are of the first author (SST), except after
a semicolon, where they are those of the fourth author (FJAD). This sequence is followed by the phytogeographic distribution class.
Alectoria nigricans (Ach.) Nyl., Empetrum nigrum heaths,
windswept knoll, on Rhododendron camtschaticum
over rock outcrop, 93–170 m, 06-32, 07-55, 13-57, 1467;070796-004.1. A.
Alectoria ochroleuca (Hoffm.) A. Massal., on rock and
gravelly Empetrum nigrum tundra in along mountain
ridge, 80–300 m, 040, 062A, 099, S1-1; 0796-005.1. A.
Allantoparmelia alpicola (Th. Fr.) Essl., on boulder in Salix arctica meadow, 230 m, 402. A.
Anaptychia setifera Räsänen, on talus at base of talus
slope, 113 m, 169W. A, B.
Arthrorhaphis alpina (Schaerer) R. Sant., meadow above
sea cliffs; 060796-491.1. A.
Asahinea chrysantha (Tuck.) Culb. & C. Culb., Empetrum
nigrum-Cladonia heath on mountain ridge, on mosses
over boulder, 200–280 m, 24-31, 344. A, M.
Bacidia sp., on coastal rock, 2 m, 153. U.
Baeomyces carneus (Retz.) Flörke, on earth in Empetrum
nigrum-lichen heath; 070796-492.1. A.
Baeomyces rufus (Hudson) Rebent., in moist rock crevices, 200 m; 140796-022.1. B.
Brigantiaea fuscolutea (Dickson) R. Sant., on mosses at cliff
base and over boulder face, 300–370 m, 376, 387. A.
Brodoa oroarctica (Krog) Goward, on boulder in Salix
barclayi thicket, 60 m, 142. A.
2002]
TALBOT ET AL.: SIMEONOF WILDERNESS LICHENS
Bryocaulon divergens (Ach.) Kärnefelt, Empetrum nigrum-moss and Empetrum nigrum-Cladonia tundra of
upper mountain slope, boulder, 225–410 m, 105, 207,
329, 23-39, 24-32, 26-43, 27-43. B.
Bryoria capillaris (Ach.) Brodo & D. Hawksw., on boulder in Heracleum maximum-Chamerion angustifolium
meadow, 100–170 m, 322; 070796-493.1. B.
Bryoria nitidula (Th. Fr.) Brodo & D. Hawksw., on mineral soil and Empetrum nigrum twigs, Empetrum nigrum-Cladina heath, on boulders and rock cliffs, 170–
300 m, 041, 064A, 07-39, 13-47, 30-32, 319, 361;
0707796-027.1, 0707796-027.2, 140796-027.3. A.
Bryoria tenuis (E. Dahl) Brodo & D. Hawksw., Empetrum
nigrum dwarf willow heath, on mosses over boulder in
Salix barclayi thicket, 60–230 m, 14-53, 149, 404. B, C.
Caloplaca scopularis (Nyl.) Lettau, on coastal rock, 2 m,
312A. A, B, C.
Cetraria aculeata (Schreber) Fr., Empetrum nigrum-dwarf
willow heath, on mosses over rock outcrops, 113–300
m, 14-64, 175, 374. A.
Cetraria ericetorum Opiz, Salix arctica-feathermoss hummock in wet gravelly fen, Empetrum nigrum-Cladina
and Empetrum nigrum-moss heath, on mosses, 20–440
m, 005, 06-56, 07-38, 107A, 112, 123, 213, 341A, 398,
13–46, 14–39, 23–35, 24–35, 24–38, 25–40, 26–25,
26–31, 27–22. A, B.
Cetraria islandica (L.) Ach., Empetrum nigrum-Salix arctica heath of upper mountain slope, 250 m, 119;
160996–068.1. A, B.
Cetraria laevigata Rass., beneath boulder on upper mountain slope, Empetrum nigrum heath, 400 m, 271;
070796–495.1, 130796–170.8, 0796–495.3. A, B, M.
Cetraria muricata (Ach.) Eckfeldt, on mosses in Anemone
narcissiflora meadow, Empetrum nigrum-Arctostaphylos uva-ursi heath, Empetrum nigrum-moss heath, on
mosses over boulder, epiphytic on Alnus viridis, 45–407
m, 071, 405, 13–55, 23–43, 25–39, 26–46, S15–8;
140796–071.1, 190796–071.2, 160796–071.3, 070796–
071.4, 120796–071.5. C.
Cetraria nigricans (Retz.) Nyl., on boulder ridge in dwarf
willow-moss tundra, Empetrum nigrum-Arctostaphylos
uva-ursi heath, 225–260 m, 134, 23–36. A.
Cetrariella delisei (Schaerer) Kärnefelt & Thell, on rock
in Empetrum nigrum heath, 400 m; 200796–066.1. A.
Cetrelia alaskana (Culb. & C. Culb.) Culb. & C. Culb.,
on boulder on mountain ridge, 250–300 m, 042, 318.
A, M.
Cetrelia cetrarioides (Duby) Culb. & C. Culb., on mosses
under cliff overhang below mountain ridge on southfacing slope, 250 m, 072. B (disjunctly circumpolar;
montane, Europe, E. Asia, Appalachians).
Cladina arbuscula (Wallr.) Hale & Culb., on mosses over
boulder in Salix arctica-Empetrum nigrum heath, 200–
477 m, 349A; 140796–083.1, 110796–083.2, 160796–
083.3, 0796–082.4. A, B.
Cladina mitis (Sandst.) Hustich, on mosses in Empetrum
nigrum-Cladina heath, Betula nana-Sphagnum poor fen
10–312 m, 003, 003A, 004, 009, 010, 012, 107, 01–25,
02–45, 03–45, 06–57, 21–33, 30–19. A, B, W.
Cladina portentosa (Dufour) Follmann subsp. pacifica
(Ahti) Ahti, Empetrum nigrum-Arctostaphylos uva-ursi
heath, 200 m, 30–29; 110796–497.1, 090796–497.2,
060797–497.3, 0796–497.4, 0796–497.5, 0796–497.6.
C (NW North America).
Cladina rangiferina (L.) Nyl., on mosses in Betula nana
poor fen, Empetrum nigrum-Vaccinium vitis-idaea
heath, on mosses over boulder, 14–250 m, 183, 342,
6017, S3–1, S3–2; 060796–164.1, 190796–164.2,
115
110796–164.3, 060796–164.4, 090796–164.5, 0796–
164.6. A, B, W.
Cladina stellaris (Opiz) Pouz. & Vezda, Empetrum nigrum heath, bog, 20–400 m; 100796–172.1, 070796–
172.2, 190796–172.3. A, B.
Cladonia amaurocraea (Flörke) Schaerer, on mosses over
cliff face, wind-exposed heath, 230–300 m, 131, 369;
190796–080.1, 120796–080.2, 070796–080.3, 0796–
080.4. A, B.
Cladonia bellidiflora (Ach.) Schaerer, on mosses over
boulder in Rubus spectabilis thicket and Rhododendron
camtschaticum heath, Empetrum nigrum heath, 15–330
m, 075, 091, 092, 109A, 125, 179, 6004, S3–5; 070796–
088.4, 180796–088.5, 200796–088.6, 0796–088.7. A,
B, C.
Cladonia borealis S. Stenroos, on mosses in Empetrum
nigrum-Arctostaphylos uva-ursi heath, Rubus spectabilis meadow, on mossy rocky, 50–250 m, 066, 073,
199, 279, 339; 140796–089.1, 070796–089.2, 070796–
089.3, 120796–089.4, 140796–089.5, 160796–089.6,
0796–089.7. A, B, W.
Cladonia cariosa (Ach.) Sprengel, on rock along coast,
meadow above sea cliffs, 5 m; 060796–093.1, 240796–
093.2. A, B, W.
Cladonia carneola (Fr.) Fr., on spruce driftwood along
coast, 2 m; 0796–094.1. A, B, W.
Cladonia chlorophaea (Sommerf.) Sprengel s.str., meadow above sea cliffs, on driftwood logs and wood of old
ranch building, coastal rocks in epilittoral zone, 2–200
m; 060796–098.1, 120796–098.2, 160796–098.3,
060796–098.4, 180796–098.5. W.
Cladonia coccifera (L.) Willd., Empetrum nigrum-Cladina hummocky heath, Empetrum nigrum-Arctostaphylos
uva-ursi heath, 30–170 m, 02–31, 06–69, 07–51, 10–
35. A, B, W.
Cladonia coniocraea (Flörke) Sprengel, Empetrum nigrum heath, 100 m; 140796–112.1. B.
Cladonia cornuta (L.) Hoffm. subsp. cornuta, on earth
between rocks, on sandy sea cliff, 20–140 m; 140796–
115.2, 070796–115.3. A, B, W.
Cladonia cornuta subsp. groenlandica (E. Dahl) Ahti, on
earth in middle mountain slope, Empetrum nigrum
heath, 30–200 m; 070796–117.1, 100796–117.2. A, B.
Cladonia crispata (Ach.) Flotow, Empetrum nigrum-Cladina hummocky heath, 30 m, 02–46. W.
Cladonia crispata (Ach.) Flotow subsp. cetrariformis (Delise) Vainio, Empetrum nigrum heath, on earth below
summit, 150–400 m; 190796–119.2, 120796–119.1,
0796–119.3. A, B.
Cladonia cyanipes (Sommerf.) Nyl., Empetrum nigrum
heath, on moss hummock in Betula nana poor fen, 5–
25 m, 087, 184. A, B.
Cladonia decorticata (Flörke) Sprengel, on rock in Empetrum nigrum heath, 120 m, 14–81. B.
Cladonia deformis (L.) Hoffm., on mosses over moist
rock of cliff face, on mosses in poor fen, Empetrum
nigrum heath, coastal driftwood log, 2–230 m, 125,
180, 182, 2046, 13–62. A, B, W.
Cladonia ecmocyna Leighton, Empetrum nigrum-Arctostaphylos uva-ursi hummocky heath, on mosses in Empetrum nigrum-dwarf willow heath, 30–120 m, 11–37,
14–83; 070796–128.1, 0796–128.2. A, B, W.
Cladonia fimbriata (L.) Fr., Empetrum nigrum-Cladina
hummocky heath, on mosses at cliff base, on mosses
over rock, Leymus mollis sandy beach, 5–300 m, 086,
368, 02–29, 22–36, S3–4; 060796–129.1, 070796–
129.1, 100796–129.3, 140796–129.4. A, B, W.
Cladonia furcata (Hudson) Schrader, Empetrum nigrum
116
THE BRYOLOGIST
heath, 70–300 m; 140796–134.1, 140796–129.4, 0796–
136.3, 0796–134.4. B.
Cladonia gracilis (L.) Willd. subsp. gracilis, Empetrum
nigrum heath, on mosses over boulder and over mosses
in Empetrum nigrum-dwarf willow heath, 5–400 m,
038, 089, 401, 06–77, 09–58, 10–22, 13–44, 13–45,
20–26, S3–3. A, B, W.
Cladonia gracilis subsp. vulnerata Ahti, over mosses in
dwarf willow-Empetrum nigrum heath of upper mountain slope, 250 m, 117. A, B.
Cladonia luteoalba Wheldon & A. Wilson, on earth, 200
m, 190796–494.1. B (interrupted circumpolar).
Cladonia maxima (Asah.) Ahti, Empetrum nigrum-Chamerion angustifolium hummocky heath, on mosses over
boulder, Calamagrostis canadensis-Salix arctica meadow, 10–200 m, 176, 185, 01–27, 02–30, 05–33, 349,
01–34, 05–41, 30–35; 070796–136.1, 0796–136.3,
070796–496.4, 0796–496.5, 0796–496.6, 140796–
496.1, 120796–496.2, 140796–496.3, 160796–136.2.
A, B, C.
Cladonia merochlorophaea var. novochlorophaea Sipman, Empetrum nigrum heath, 200 m; 120796–150.1.
A, B.
Cladonia pleurota (Flörke) Schaerer, on mosses over
moist rock of cliff face, 230 m, 127A. A, B, W.
Cladonia phyllophora Hoffm., on mosses over rocks, 50–
100 m; 160796–155.1, 100796–155.2. A, B.
Cladonia pocillum (Ach.) Grognot, epiphytic on base of
Alnus viridis trunk, 50 m, 081. A.
Cladonia pyxidata (L.) Hoffm., on earth in Empetrum nigrum heath, 20 m; 160796–162.1. A, B, W.
Cladonia ramulosa (With.) J. R. Laundon, Empetrum nigrum heath, on earth near sea, 5–200 m; 190796–163.1,
140796–163.2. B, T.
Cladonia scabriuscula (Delise) Nyl., on mosses over
boulder in Salix arctica-Empetrum nigrum heath, 200
m, 347. B, C, W.
Cladonia squamosa Hoffm., Empetrum nigrum-Cladina
hummocky heath, on mosses over rocks, Calamagrostis
canadensis-Salix arctica meadow, 30–300 m, 276, 338,
371, 380, 6017A, 02–39, 09–53; 160796–170.1,
060796–170.2, 070796–170.3, 140796–170.4, 120796–
170.5, 140796–170.6, 060796–170.7, 130796–170.8,
0796–170.9. A, B, W.
Cladonia stricta (Nyl.) Nyl., Empetrum nigrum-Cladina/
moss hummocky heath, on rock in Empetrum nigrumdwarf willow heath, on humus, 30–170 m, 084, 02–37,
10–37, 14–80, 20–25, 21–23. A.
Cladonia subfurcata (Nyl.) Arnold, Empetrum nigrumCladina heath, on moss hummock in Betula nana poor
fen, 25–93 m, 184A, 06–68. A, B, C.
Cladonia uncialis (L.) F. H. Wigg., Empetrum nigrum
heath, Salix arctica-moss poor fen, 20–200 m, 013;
060796–187.1, 070796–187.2, 190796–187.3, 0796–
187.4, 0796–187.5, 120796–186.1. A, B, W.
Collema tenax (Sw.) Ach., Empetrum nigrum-Salix arctica hummocky heath, on mosses at cliff base, 107–300
m, 372, 09–41. A, B, W.
Cornicularia divergens Ach., Empetrum nigrum heath,
100 m; 0796–197.1. A.
Ephebe lanata (L.) Vainio, epiphytic on moss hummock,
150 m, 6023. A.
Flavocetraria cucullata (Bellardi) Kärnefelt, Salix arctica-feathermoss hummock in wet gravelly fen, Empetrum nigrum-moss heath, on mosses over boulder, 20–
200 m, 341B, 006, 12–34; 060796–065.1, 160796–
065.2. A, B.
Fulgensia bracteata (Hoffm.) Räsänen, on wooden ranch
[VOL. 105
house shingles surrounded by Empetrum nigrum-Salix
arctica hummocky heath, 6 m, 268B. A, B, W.
Fuscopannaria praetermissa (Nyl.) P. M. Jørg., on moist
cliffs, 100 m; 070796–001B.1. A, B.
Hypogymnia physodes (L.) Nyl., over mosses on rocks,
400 m; 140796–225.2. W.
Hypogymnia subobscura (Vainio) Poelt, on mosses over
rock in Salix arctica-Rhododendron camtschaticum
heath, 250 m, 032. A.
Hypogymnia vittata (Ach.) Parrique, moist rock of cliff
face, on mosses over rocks, 230–300 m, 122, 363;
160796–228.1, 160796–228.2. A.
Lecanora beringii Nyl. (‘‘behringii’’), on coastal boulder
in splash zone, 2 m, 160. A.
Lecanora epibryon (Ach.) Ach., over mosses and Empetrum nigrum in Empetrum nigrum heath of upper mountain slope, 330 m, 111. A.
Lecanora orae-frigidae R. Sant., xylicolous on wooden
fence, 13 m, 137, 138. A, B.
Lecanora symmicta (Ach.) Ach., epiphytic on Alnus viridis
on south-facing lower mountain slope, 50 m, 079. B.
Lecidea lapicida (Ach.) Ach., on boulder on mountain
ridge and Angelica lucida-Carex macrochaeta meadow,
on stones in gravelly tundra of upper mountain slope,
170–400 m, 046, 108, 209A, 330, 07–80X. A, B, C.
Lecidea plebeja Nyl., on wooden ranch house shingles
surrounded by Empetrum nigrum-Salix arctica hummocky heath, 6 m, 268C. B (NA & Europe).
Lecidella anomaloides (A. Massal.) Hertel & R. Kilias (5
Lecidea goniophila auct.), on boulder on mountain
ridge, 250 m, 050. U.
Lecidella stigmatea (Ach.) Hertel & Leuckert, epilithic on
boulder mid-slope mountain canyon, 170 m, 6030. A.
Leptogium corniculatum (Hoffm.) Minks, on mosses over
wind protected rock in Geranium erianthum meadow,
on rock outcrop, on mosses over moist shady rock of
cliff face, 113–370 m, 024, 128, 171, 203; 070796–
269.1. A & B (w NA & w & e Eurasia), C.
Lobaria cf. isidiosa (Müll. Arg) Vainio, epiphytic on Alnus viridis, 40 m, S15–6. U (Not in NA).
Lobaria hallii (Tuck.) Zahlbr., epiphytic on Alnus viridis
in Alnus viridis thicket, 40–66 m, S10–2, S15–2, S15–4,
S15–5; 140796–498.1. C (disjunct: w NA & Greenland)
Lobaria kurokawae Yoshim., on mosses over moist shaded boulder in Carex meadow, 370 m, 206. M.
Lobaria linita (Ach.) Rabenh., Empetrum nigrum-moss
heath from sea level to upper mountain slopes, on
mosses over coastal cliff, 2–280 m, 021, 313, 334, 357,
24–44, S3–7; 120796–499.1, 190796–499.2, 140796–
499.3, 060796–499.4, 070796–499.5, 160796–499.6,
0796–499.7. A, C.
Lobaria scrobicularia (Scop.) DC., on willow, 70 m;
0796–278.4. B, C.
Melanelia elegantula (Zahlbr.) Essl., epiphytic on Alnus
viridis in Alnus viridis thicket, 100 m, 190. A, B.
Melanelia hepatizon (Ach.) Thell, on rock in Empetrum
nigrum-moss tundra, Geranium erianthum meadow, Salix barclayi, and Angelica lucida-Carex macrochaeta
meadow, 60–310 m, 030, 106, 135, 145, 146, 178, 328.
A, B, W.
Melanelia stygia (L.) Essl., on boulder on stone-stripe
mountain ridge, 250 m, 056, 059. A, B.
Mycoblastus alpinus (Schaerer) Schauer, on boulder on
mountain ridge in Arctostaphylos alpina-Vaccinium vitis-idaea heath, 200–250 m, 049, 332. A, B. Reported
by Rothrock (1884) as Heterothecium sanguinarium
(Fl.) Tuck. var. alpinum Fr. for the Shumagin Islands.
Nephroma arcticum (L.) Torss., Loiseleuria procumbens-
2002]
TALBOT ET AL.: SIMEONOF WILDERNESS LICHENS
Carex heath, on boulder, Carex-moss meadow, 300–
400 m, 186, 320, 395; 140796–288.1. A, B.
Nephroma parile (Ach.) Ach., epiphytic on alder, Empetrum nigrum-Salix arctica hummocky heath 20–50 m
083, 09–39. B.
Ochrolechia frigida (Sw.) Lynge, Empetrum nigrum-Cladina hummocky heath, on boulder on mountain ridge,
moss-Empetrum nigrum tundra, on mosses in Empetrum
nigrum heath, 20–440 m, 008, 052, 057, 058, 063A,
064B, 208, 03–46, 06–64, 09–37, 25–45, 27–23;
070796–296.1, 070796–296.2, 060796–296.3, 190796–
296.4, 140796–296.5, 190796–296.6, 060796–296.7. A.
Ochrolechia frigida fo. gonatodes (Ach.) Lynge, epilithic
on boulder in Empetrum nigrum heath, 200 m, 340. A.
Ochrolechia subplicans (Nyl.) Brodo, on coastal boulder
in splash zone, 2 m, 164. A.
Ochrolechia tartarea (L.) A. Massal., on boulder in Salix
arctica-Empetrum nigrum heath, 200 m, 345B. A, B,
C. Reported by Cummings (1910) as Lecanora tartarea
(L.) Ach. for the Shumagin Islands.
Ochrolechia upsaliensis (L.) A. Massal., on humus over
boulder, 300 m, 316. A.
Ochrolechia xanthostoma (Sommerf.) K. Schmitz &
Lumbsch, epiphytic on Salix glauca and Alnus viridis,
20–50 m, 017, 077, 082. A, B, C.
Ophioparma lapponica (Räsänen) Hafellner & R. W. Rogers, on boulder on mountain ridge and in Arctostaphylos alpina-Vaccinium vitis-idaea heath, 200–250 m,
051, 331, 353. A.
Omphalina hudsoniana (H. S. Jenn.) H. E. Bigelow, on soil
on hillside, 200 m; 1140796–500.220796–500.1,. B.
Pannaria conoplea (Ach.) Bory, on mosses and lichens
over boulder in Salix arctica-Empetrum nigrum heath,
200 m, 350. B.
Pannaria pezizoides (Weber) Trevisan, Empetrum nigrumCladina heath, on mosses over rock face in cliff chimney, on soil in meadow above sea cliffs, 93–300 m, 06–
58, 365; 060796–306.1. A, B, C.
Parmelia fraudans (Nyl.) Nyl., epiphytic on Alnus viridis
in Alnus viridis thicket and on introduced Picea sitchensis, 100–265 m, 191, 265. A, B.
Parmelia omphalodes (L.) Ach., on rock in Empetrum nigrum-Salix arctica heath, on wind-protected boulder on
mountain ridge, talus, Empetrum nigrum-Arctostaphylos alpina heath, epiphytic on Alnus viridis, Rhododendron camtschaticum, and mosses, 47–300 m, 036, 048,
053, 116, 169, 177, 23–58, 270, 272, 346, 375, 381;
140796–323.1. A, B.
Parmelia saxatilis (L.) Ach., on rock outcrop in mid-slope
mountain canyon wall, boulders in Salix barclayi thicket, Carex meadows, Heracleum maximum-Chamerion
angustifolium meadow, Salix arctica-Carex meadows,
epiphytic on Alnus viridis, epilittoral rocks, on old
wood of ranch house, 6–370 m, 097, 143, 170, 173,
202, 269, 317, 326, 406A, S15–1, S9–1; 060796–326.1,
180796–326.2, 0796–326.3. A, B, W.
Parmelia squarrosa Hale, on boulder in Geranium erianthum meadow, 250 m, 029. B.
Parmelia sulcata Taylor, epiphytic on Salix glauca, Alnus
viridis, and introduced Picea sitchensis, on boulder in
Geranium erianthum meadow and on mosses over
boulder, coastal driftwood log, Empetrum nigrum heath,
Rubus spectabilis-Calamagrostis canadensis meadow,
on wood of old ranch house, 2–250 m, 019, 027, 078,
165, 196, 253, 257, 267, 337, B3, C1, S10–3, S15–3;
120796–331.4, 180796–331.5. A, B, W.
Peltigera aphthosa (L.) Willd., Empetrum nigrum heaths,
Alnus viridis thickets, on mosses, 12–370 m, 02–16,
04–30, 05–42, 07–53, 08–28, 09–35, 090, 095, 11–28,
117
12–31, 13–38, 168W, 21–29, 23–42, 25–43, 29–26,
S10–1, S3–8; 070796–335.1, 120796–335.2, 140796–
335.3, 100796–335.4, 0796–335.5. Reported by Thomson (1984) for the Shumagin Islands. A, B, W.
Peltigera britannica (Gyelnik) Holt-Hartw. & Tønsberg,
on mossy boulder in Heracleum maximum-Chamerion
angustifolium meadow, Empetrum nigrum heath, on
mosses over rock face in cliff chimney, 100–300 m,
323, 335, 362, 366A, 370. C, M.
Peltigera canina (L.) Willd., Empetrum nigrum heaths, on
cliffs near sea, on mosses, 30–225 m, 06–78, 14–68,
21–25, 22–37, 23–57; 070796–336.1, 200796–336.3,
160796–336.4, 060796–339.5. A, B, W.
Peltigera collina (Ach.) Schrader, epiphytic on Salix glauca, Alnus viridis, and Sambucus racemosa, Empetrum
nigrum-Salix arctica heath, coastal driftwood log, 20–
200 m, 016, 020, 080, 09–46, 09–56, 174, 359, 2045,
6021. A, B.
Peltigera didactyla (With.) J. R. Laundon, Leymus mollis
sandy bench, 5 m, 085. A, B, W.
Peltigera leucophlebia (Nyl.) Gyelnik, on mosses in meadows, 113–250 m 070, 167; 140796–341.1. A, B, W.
Peltigera malacea (Ach.) Funck, Empetrum nigrum-Arctostaphylos uva-ursi hummocky heath in depression, 30
m, 11–80. A, B, W.
Peltigera membranacea (Ach.) Nyl., Empetrum nigrum
heaths, on mosses over wind-protected rock, epiphytic
on Sambucus racemosa, 5–370 m, 01–33, 023, 04–26,
05–35, 08–19, 09–23, 10–38, 11–41, 12–32, 13–35,
159, 201, 28–27, 29–29, 30–22, 30–34A, 321, 367,
6020, S3–6, 069, 074; 0796–343.1, 0796–343.2, 0796–
343.3. B, C, W.
Peltigera neckeri Müll. Arg., on cliffs along the coast,
Leymus mollis meadow, 4 m; 060796–002B.1, 240796–
002B.2. B.
Peltigera polydactylon (Necker) Hoffm., Empetrum nigrum heaths, on mosses over moist cliff, 225–440 m,
23–56, 24–45, 27–53, 378, 13–63. B, C, W.
Peltigera rufescens (Weiss) Humb., Rubus spectabilisCalamagrostis canadensis thicket, Leymus mollis
meadow, 2–40 m, B1; 240796–346.1. A, B.
Peltigera scabrosa Th. Fr., on mosses over moist rock of
cliff face, Empetrum nigrum heaths, 170–370 m, 093,
121, 124, 126, 129, 131A, 25–49, 25–51, 366; 140796–
347.1, 100796–347.2, 120796–347.3. A, B, C, W.
Peltigera scabrosella Holt.-Hartw., on mosses, 65 m;
140796–003B.1. A, B (interrupted circumpolar).
Peltigera venosa (L.) Hoffm., on eroding sea bluffs on
mineral soil and mosses, 4 m, 273. A, B.
Pertusaria alaskensis Erichsen, on humus over boulder,
300 m, 315. M.
Pertusaria bryontha (Ach.) Nyl., Empetrum nigrum-Cladina heath, 170 m, 07–58. A.
Pertusaria coriacea (Th. Fr.) Th. Fr., on boulder in Empetrum nigrum-moss tundra, Empetrum nigrum-dwarf
willow heath, 120–310 m, 106A, 14–65, 345A. A.
Pertusaria dactylina (Ach.) Nyl., Empetrum nigrum-Cladina heath, 93 m, 06–71. A.
Pertusaria panyrga (Ach.) A. Massal., epiphytic on Salix
glauca, on rock, 20–53 m, 018, 197A. A.
Pertusaria subobducens Nyl., epiphytic on Alnus viridis,
100–200 m, 187, 358. A.
Physcia adscendens (Fr.) H. Olivier, on introduced Picea
sitchensis surrounded by Empetrum nigrum-Salix arctica hummocky heath 6 m, 259, 263. A, B, W.
Physcia caesia (Hoffm.) Furnr., on rocks, gravestone of
John C. Smith (1815–1917), epiphytic on Picea sitchensis, on wooden ranch house, on driftwood, 2–300 m,
118
THE BRYOLOGIST
136, 152, 154, 162, 252, 254, 262, 266, 311, 314, 336;
120796–370.1. A, B, W.
Physconia muscigena (Ach.) Poelt var. muscigena, meadow above sea cliffs, 15 m; 060796–004B.1. A, B, W.
Pilophorus acicularis (Ach.) Th. Fr., on rocks in Geranium erianthum meadow, Empetrum nigrum heaths,
Rhododendron camtschaticum-moss heath, Calamagrostis canadensis-Salix arctica meadow, 93–250 m,
022, 06–90X, 118, 118A, 274, 6003, 6018. C, M.
Pilophorus robustus Th. Fr., on stones and rocks, 200 m;
160796–005B.1, 160796–005B.2, 140796–005B.3. A.
Placopsis gelida (L.) Lindsay, over rock in Empetrum nigrum-Carex heath of upper mountain slope, on soil, 300
m, 115; 180796–384.1. A, B, C, W.
Platismatia glauca (L.) Culb. & C. Culb., on introduced
Picea sitchensis, on boulders near summit, 6–410 m,
261; 190796–388.1. B.
Platismatia lacunosa (Ach.) Culb. & C. Culb., epiphytic
on Alnus viridis, on rock in Empetrum nigrum heath,
33–100 m, 195, S17–2; 190796–006B.1, 070796–
006B.2, 060796–006B.3, 060796–006B.4. C (NA endemic).
Porpidia flavocaerulescens (Hornem.) Hertel & A. J.
Schwab, on rock in Empetrum nigrum heaths along
mountain ridge, 250–370 m, 063, 114, 383. A.
Porpidia grisea Gowan, epilithic on stones and boulders
in Empetrum nigrum gravelly heath of upper-mountain
slope, also on boulder in Arctostaphylos alpina-Vaccinium vitis-idaea heath, 200–330 m, 110, 333. A, B.
Pseudephebe pubescens (L.) Choisy, on boulder in Geranium erianthum meadow and mountain ridges, also
on rock in Eriophorum angustifolium peatland, 3–370
m, 026, 047, 382, S5–1; 190796–395.1, 070796–395.2.
A, B.
Pseudocyphellaria anomala Brodo & Ahti, epiphytic on
Alnus viridis, on rock in Empetrum nigrum-Arctostaphylos uva-ursi hummocky heath, 12–100 m, 076, 189B,
193, 277; 190796–008B.1, 190796–008B.2. C (w NA
endemic).
Pseudocyphellaria anthraspis (Ach.) H. Magn., on mosses
over rock in Salix arctica-Rhododenron camtschaticum
heath, on mosses in Empetrum nigrum heath, 250 m,
033, 037, 067. C (w NA endemic).
Pseudocyphellaria crocata (L.) Vainio, epiphytic on Alnus
viridis, 40–100 m, 189A, S15–7; 190796–007B.1. B
(incompletely circumpolar).
Psoroma hypnorum (Vahl.) Gray, Empetrum nigrum-Arctostaphylos uva-ursi/Cladina heath, epiphytic on Salix
glauca, 45 m, 13–64, 360; 140796–009B.1. A, B, C.
Ramalina almquistii Vainio, on rocks and boulders in
heaths, meadows, and ridges, 80–250 m, 044, 034, 324,
S1–2. M.
Ramalina scoparia Vainio, xylicolous on wooden fence,
13 m, 140. M.
Rhizocarpon anseris Lynge, on rock on mountain ridge in
Loiseleuria procumbens-moss heath, 400 m, 209B. A.
Rhizocarpon disporum (Hepp) Müll. Arg., on boulder in
Heracleum maximum-Chamerion angustifolium meadow, on rock cliff, 100–300 m, 325, 370A. A.
Rhizocarpon eupetraeoides (Nyl.) Blomb. & Forss., boulder face, 370 m, 385. A.
Rhizocarpon geographicum (L.) DC., on boulders, 200–
250 m, 045, 352. A, B, W.
Rhizocarpon obscuratum (Ach.) A. Massal., boulder face,
370 m, 386. A, B, C.
Rhizocarpon polycarpum (Hepp) Th. Fr., on rock in Carex-moss meadow, 400 m, 397. A, B.
Siphula ceratites (Wahlenb.) Fr., on rocks in Arctostaphylos
alpina-Vaccinium vitis-idaea heath, 200 m, 331A. A.
[VOL. 105
Solorina crocea (L.) Ach., Empetrum nigrum gravelly tundra of upper mountain slope, on mineral soil on coastal
wind-eroded terrace, 8–300 m, 104, 275. A, C.
Sphaerophorus fragilis (L.) Pers., Empetrum nigrum-Salix
arctica hummocky heath, on rocks in heaths and Eriophorum angustifolium peatland, 3–280 m, 09–57, 24–
80, S5–2; 070796–101B.1. A.
Sphaerophorus globosus (Hudson) Vainio, on hummock
in wet gravelly fen, on boulder on mountain ridge, 20–
250 m, 011, 043; 070796–423.1, 060796–423.2,
140796–423.3. Reported by Cummings (1910) and
Rothrock (1884) for the Shumagin Islands (Little Koniuji Island). A.
Stereocaulon alpinum Funck, Empetrum nigrum tundra of
upper mountain slope, on mosses in Fragaria chiloensis-Empetrum nigrum meadow, 5–00 m, 102, 158. A.
Stereocaulon arenarium (Savicz) Lamb, on rock outcrop,
boulder on mountain ridge, on rock in gravelly fen
streamlet, on mosses over boulder, 20–400 m, 001, 007,
028, 035, 039, 055, 060, 062, 094, 098, 103, 109, 198,
210, 351, 354, 384, 389, 394, 408. M.
Stereocaulon coniophyllum Lamb, on rock in Empetrum
nigrum heath, 30 m, S9–3. A.
Stereocaulon grande (H. Magn.) H. Magn., on mosses in
Empetrum nigrum-moss heath, on gravelly tundra of
upper mountain slope, 250–300 m, 031, 101. A, B.
Stereocaulon intermedium (Savicz) H. Magn., on mosses
over cliff face, on rock in Empetrum nigrum heath,
113–230 m, 127, 130, 14–82, 172, 355. C, M.
Stereocaulon paschale (L.) Hoffm., Empetrum nigrum
heaths, on mosses in Empetrum nigrum heaths, Carex
macrochaeta-moss meadow, 14–440 m, 06–54, 07–46,
09–36, 12–48, 13–23, 14–34, 214, 23–41, 24–21, 25–
56, 27–30, 396, 400, S3–9; 0796–437.1, 0796–437.2.
A, B, W.
Stereocaulon saxatile H. Magn., on boulder in Angelica
lucida-Carex macrochaeta meadow, 100 m, 327. A, B.
Stereocaulon subcoralloides (Nyl.) Nyl., over rock in Empetrum nigrum heath of upper mountain slope, on
mosses over rock face in cliff chimney, 300–330 m,
113, 364. B.
Stereocaulon tomentosum Fr., on mosses in Empetrum nigrum-Arctostaphylos uva-ursi heath below mountain
ridge, 250 m, 068. A, B.
Sticta arctica Degel., Empetrum nigrum heath, on mosses
and liverworts over boulder and moist cliffs, 113–300
m, 07–40, 133, 167W, 204, 343, 345, 346A, 348, 356,
377; 070796–011B.1, 070796–011B.2. M.
Sticta limbata (Sm.) Ach., epiphytic on Salix glauca, 20
m, 014, 015. W.
Sticta weigelii (Ach.) Vainio, on mosses over boulder on
lower mountain slope, 50 m, 083B; 120796–012B.1,
0796–12B.2. W (Tropical to Temperate, disjunct in
Alaska).
Thamnolia subuliformis (Ehrh.) Culb., Empetrum nigrum
heath along mountain ridge, 250 m, 061, 064, 065;
140796–453.1, 0796–453.2. A.
Thamnolia vermicularis (Sw.) Schaerer, Empetrum nigrum
hummocky heaths, on pebbles and rocks on mountain
ridge, 33–440 m, 03–29, 06–30, 07–30, 09–27, 13–29,
14–28, 16–30, 211, 212, 23–20, 24–17, 25–29, 26–13,
27–17, S17–1; 140796–454.1, 0796–454.2. A.
Trapeliopsis granulosa (Hoffm.) Lumbsch, humus on Empetrum nigrum heath hummock, 5 m, 088. A, B.
Tuckermannopsis chlorophylla (Willd.) Hale, on mosses
over boulder in Salix barclayi thicket, on coastal rock,
2–370 m, 148, 205, 302; 040796–063.1. A.
Tuckermannopsis platyphylla (Tuck.) Hale, epiphytic on
Alnus viridis, 100 m, 192. B, T (w NA).
2002]
TALBOT ET AL.: SIMEONOF WILDERNESS LICHENS
119
TABLE 1. Frequently occurring lichens in the crowberry (Empetrum nigrum) heaths of Simeonof Island, Alaska,
based on their presence in 30 relevés. To be included a species must occur in at least two relevés. Key to presence
classes: I, 1–20%; II, 21–40%; III 5 41–60%; IV, 61–80%; V, 81–100%.
Cladina rangiferina
Cladina mitis
Cladonia gracilis
Lobaria linita
Peltigera membranacea
Peltigera aphthosa
Cladonia maxima
Cladina stellaris
Cladonia uncialis
Thamnolia vermicularis
Flavocetraria cucullata
Sphaerophorus globosus
IV
IV
IV
IV
III
III
III
III
III
III
II
II
Cladonia amaurocraea
Ochrolechia frigida
Cetraria ericetorum
Alectoria nigricans
Alectoria ochroleuca
Cladonia crispata
Stereocaulon paschale
Cladonia bellidiflora
Peltigera canina
Cladina cf. stellaris
Cladonia coccifera
Cladonia squamosa
II
II
II
II
II
II
II
II
II
II
II
II
Species occurring with a presence of less than 20%: Bryocaulon divergens, Bryoria nitidula, Cladina pacifica,
Cladonia chlorophaea, C. deformis, C. fimbriata, C. furcata, C. stricta, Cetraria muricata, Parmelia omphlodes, Peltigera polydactylon, P. scabrosa, Pertusaria dactylina, Psoroma hypnorum, Sticta arctica, S. weigelii, and Thamnolia
subuliformis.
Umbilicaria angulata Tuck., chimney cliff, 200 m, 280.
A (w NA endemic).
Umbilicaria arctica (Ach.) Nyl., on boulder in Arctostaphylos alpina-Vaccinium vitis-idaea heath, 200 m,
332A. A.
Umbilicaria havaasii Llano, on boulders in Cladina lichen
meadow and Salix arctica-Carex macrochaeta meadow, 53–400 m, 197, 399, 403, 406, 407. A (amphiAtlantic species with broad disjunctions).
Umbilicaria hyperborea (Ach.) Hoffm., on mosses over
wind-protected rock in a Geranium erianthum meadow,
on rock in Empetrum nigrum heath, 30–250 m, 025,
S9–2; 070796–470.1. A, B, W.
Umbilicaria proboscidea (L.) Schrader, on boulder on
mountain ridge and in Salix barclayi thicket, 60–370
m, 054, 144, 392; 140796–473.1, 070796–473.2,
070796–473.3. A.
Umbilicaria scholanderi (Llano) Krog, on cliff face, 300
m, 373. M (disjunct locations in NA).
Umbilicaria torrefacta (Lightf.) Schrader, on stones, 100
m; 070796–476.2. A.
Umbilicaria vellea (L.) Hoffm., dry exposed rock of cliff
face and on wind-protected boulder, 230–370 m, 120,
200; 070796–477.1, 070796–477.2, 070796–477.3. A, B.
Verrucaria aethiobola Wahlenb., on rock in stream where
stream enters harbor, 2 m, 290A, 290B. C, W.
Verrucaria arctica Lynge, on coastal rock, 2 m, 151. A.
Verrucaria ceuthocarpa Wahlenb., on coastal rock, 2 m,
312B. A, C.
Verrucaria maura Wahlenb., on coastal rock, 2 m, 155.
C, W.
Xanthoria candelaria (L.) Th. Fr., on coastal boulder in
splash zone, on coastal driftwood log in Leymus mollis
beach meadow, on wooden ranch house, 2–12 m, 161,
166, 250, 268A, 278. A, B, W.
Xanthoria elegans (Link) Th. Fr., on coastal rock, 2 m,
150, 156, 163. A, B, W.
Xanthoria sorediata (Vainio) Poelt, xylicolous on wooden
fence, 13 m, 139. A, B.
Lichen components of the vegetation. Common
lichen species occurring in oceanic crowberry (Empetrum nigrum) heaths are shown in Table 1. The
most frequent lichens (occurring with a presence of
40% or greater) are listed in decreasing order of
presence: Cladina rangiferina, C. mitis, Cladonia
gracilis, Lobaria linita, Peltigera aphthosa, P.
membranacea, Cladina stellaris, Cladonia maxima,
C. uncialis, and Thamnolia vermicularis.
DISCUSSION
The high percentage of arctic-alpine and boreal
elements reflects the northern geographical location
of Simeonof Island. Similarly, the moderately abundant coastal element is reflected in the maritime
climate. It is of considerable interest that the proportionate representation of phytogeographic categories is similar between the sites listed for western
Alaska in Figure 3. This is also the pattern for both
Kodiak Island (Talbot 1998) and Tuxedni Wilderness Area (Talbot et al. 1992).
Our data on frequently occurring lichens in Empetrum nigrum dwarf shrub heath present basic information for comparison with other northern areas.
Comparable data from along a lowland to alpine
mesotopographic gradient on Mount Simeon in Izembek National Wildlife Refuge show close similarity in the common species present. Seventy-one
percent of the frequently occurring lichens on Izembek (Talbot et al. 2000; Table 2, community
types 5, ‘‘Empetrum nigrum-Chamerion angustifolium heath’’ and community type 6, ’’Empetrum nigrum-Loiseleuria procumbens heath’’) were also
important in the heath vegetation of Simeonof Island. Important lichens on Izembek listed in decreasing order of presence include Cladina rangiferina, C. mitis, Cladonia gracilis, Lobaria linita,
Peltigera aphthosa, P. membranacea, Cladonia uncialis, Thamnolia vermicularis, Flavocetraria cucullata, Sphaerophorus globosus, Cladonia amaurocraea, Ochrolechia frigida, Cetraria ericetorum,
Alectoria nigrescens, A. ochroleuca, Stereocaulon
paschale, Nephroma arctica, and Cladonia squamosa. Similarly, data from along a lowland to al-
120
THE BRYOLOGIST
pine mesotopographic gradient on Kodiak Island
(Talbot 1998, Table 1, community type 1, ‘‘Empetrum nigrum-Oxytropis nigrescens heath’’ and community type 2, ‘‘Empetrum nigrum-Festuca altaica
heath’’), show close similarity in the common species present, but differences in the percent presence
of each. Common lichens of western Kodiak Island
arranged in order of decreasing presence include:
Peltigera membranacea, P. aphthosa, Sphaerophorus globosus, Stereocaulon alpinum, Cladonia
gracilis, Cladina rangiferina, Lobaria linita, Cladina mitis, Cladonia coccifera, Thamnolia subuliformis, Ochrolechia frigida, Cladina stellaris, Cetraria laevigata, Cladonia uncialis, Alectoria nigricans, Peltigera malacea, and Cetraria muricata.
The observed importance of frequently occurring
lichens on Simeonof Island was consistent with
data from similar heath vegetation from Izembek
(Talbot et al. 2000) and Kodiak (Talbot 1998). Similar studies (SST) are underway to determine if this
pattern applies to the central and eastern portions
of the Alaska Peninsula.
In discussing lichen communities on heath lands,
Ahti and Oksanen (1990) noted that they contain
mainly oceanic, circumpolar species. Four of the
five taxa listed by Ahti and Oksanen (1990) as preferring oceanic heaths are common on Simeonof
Island, coinciding with their observation—Cetraria
aculeata (common in study area heaths but recorded in one relevé), Cladonia gracilis subsp. gracilis,
Ochrolechia frigida, and Sphaerophorus globosus;
a fifth, Hypogymnia physodes, was not collected.
ACKNOWLEDGMENTS
We are grateful to the Air Quality Branch, Division of
Refuges, U.S. Fish and Wildlife Service, Denver; Alaska
Maritime National Wildlife Refuge, Homer; U.S. National
Park Service, Anchorage, and the German Research Foundation (DFG) for funding support; and to the crew of the
M/V Tiglax for safe passage. Finally, we appreciate the
comments of Theodore L. Esslinger and one anonymous
reviewer.
LITERATURE CITED
AHTI, T. 1980. Taxonomic revision of Cladonia gracilis
and its allies. Annales Botanici Fennici 17: 195–243.
AHTI, T. & J. OKSANEN. 1990. Epigeic lichen communities
of taiga and tundra regions. Vegetatio 86: 39–70.
ARUP, U. 1994. The genus Caloplaca on seashore rocks
in eastern North America. THE BRYOLOGIST 97: 377–
392.
BURK, C. A. 1965. Geology of the Alaska Peninsula-island
arc and continental margin. Geological Society of
America Memoirs 99: 1–250.
CULBERSON, W. L. & C. F. CULBERSON. 1968. The lichen
genera Cetrelia and Platismatia (Parmeliaceae). Contributions to the United States National Herbarium 34:
449–558.
——— & ———. 1978. Cetrelia cetrarioides and C.
[VOL. 105
monachorum (Parmeliaceae) in the New World. THE
BRYOLOGIST 81: 517–523.
CUMMINGS, C. E. 1910. The lichens of Alaska, pp. 67–
149. In J. Cardot, C. E. Cummings, A. W. Evans, C.
W. Peck, P. A. Saccardo, D. A. Saunders, I. Theriot &
W. Trelease, Cryptogamic Botany, Harriman Alaska
Series, Vol. 5. Smithsonian Institution, Washington,
D.C.
DANIËLS, F. J. A., S. S. TALBOT, S. L. TALBOT & W. B.
SCHOFIELD. 1998. Geobotanical aspects of Simeonof
Island, Shumagin Islands, Alaska. Berichte der Reinhold-Tüxen-Gesellschaft 10: 125–138.
DIBBEN, M. J. 1980. The chemosystematics of the lichen
genus Pertusaria in North America North of Mexico.
Milwaukee Public Museum Publications in Botany
No. 5: 1–162.
ESSLINGER, T. L. 1998. A cumulative checklist for the lichen-forming, lichenicolous and allied fungi of the
continental United States and Canada. North Dakota
State University: http://www.ndsu.nodak.edu/instruct/
esslinge/chcklst/chcklst7.htm (First posted on 1 December 1997, Most recent update: 18 September
1998), Fargo, ND.
——— & R. S. EGAN. 1995. A sixth checklist of the lichen-forming, lichenicolous, and allied fungi of the
continental United States and Canada. THE BRYOLOGIST 98: 467–549.
FOX, D. G., J. C. BERNABO & B. HOOD. 1987. Guidelines
for measuring the physical, chemical, and biological
condition of wilderness ecosystems. USDA Forest
Service Technical Report RM-146. Rocky Mountain
Forest and Range Experiment Station, Fort Collins,
CO.
GEISER, L. H., K. L. DILLMAN, C. C. DERR & M. C. STENSVOLD. 1994. Lichens of southeastern Alaska. USDA,
Forest Service No. R10-TB-45. Tongass National Forest, Petersburg, AK.
GOULD, W. A. 1994. Macrolichens of the Coppermine,
Hood, and Thomsen Rivers, Northwest Territories.
THE BRYOLOGIST 97: 42–46.
GOWARD, T., B. MCCUNE & D. MEIDINGER. 1994. The Lichens of British Columbia. Illustrated Keys. Part 1—
Foliose and Squamulose Species. Special Report Series, 8. Research Program, British Columbia Ministry
of Forests, Victoria, BC.
HEUSSER, C. J. 1983. Pollen diagrams from the Shumagin
Islands and adjacent Alaska Peninsula, southwestern
Alaska. Boreas 12: 279–295.
HULTÉN, E. 1941–50. Flora of Alaska and Yukon, 1–10.
Lunds Universitets Årsskrift N. F., Avd. 2, vols. 37–
46.
JAHNS, H. M. 1981. The genus Pilophorus. Mycotaxon 13:
289–330.
KROG, H. 1968. The macrolichens of Alaska. Norsk Polarinstitutt Skrifter 144: 1–180.
LESLIE, L. D. 1989. Alaska climate summaries. Alaska
Climate Center Technical Note 5, 2nd ed. Arctic Environmental Information and Data Center, University
of Alaska, Anchorage.
RIEGER, S., D. B. SCHOEPHOSTER & C. E. FURBISH. 1979.
Exploratory Soil Survey of Alaska. USDA, Soil Conservation Service, Washington, D.C.
ROTHROCK, J. T. 1884. List of, and notes upon, the lichens
collected by Dr. T. H. Bean in Alaska and the adjacent
region in 1880. Proceedings of the United States National Museum 7: 1–9.
SEARBY, H. W. 1959. Climate of Alaska. U.S. Department
of Commerce, Environmental Data Service, Washington, D.C.
2002]
TALBOT ET AL.: SIMEONOF WILDERNESS LICHENS
SøCHTING, U. & M. OLECH. 1995. The genus Caloplaca
in polar regions. Lichenologist 27: 463–471.
SWAIN, U. G. 1990. National Natural Landmarks in Alaska. Unpublished report. Alaska Natural Heritage Program, The Nature Conservancy, Anchorage, AK.
TALBOT, S. S. 1998. Lichens of the Spiridon Peninsula,
western Kodiak Island, Alaska, with observations on
lichen communities, pp. 245–260. In M. G. Glenn, R.
C. Harris, R. Dirig & M. S. Cole (eds.), Lichenographia Thomsoniana: North American Lichenology in
Honor of J. W. Thomson. Mycotaxon Ltd., Ithaca, NY.
———, S. L. TALBOT, & J. W. THOMSON. 1991. Lichens
of Attu Island, Alaska. THE BRYOLOGIST 94: 421–426.
———, ——— & ———. 1992. Lichens of Tuxedni
Wilderness Area, Alaska. THE BRYOLOGIST 95: 20–30.
———, ———, ——— & W. B. SCHOFIELD. 1997. Lichens of Adak Island, central Aleutian Islands, Alaska.
THE BRYOLOGIST 100: 241–250.
———, ———, ——— & ———. 2000. Lichens of Izembek National Wildlife Refuge, westernmost Alaska
Peninsula. THE BRYOLOGIST 103: 379–389.
121
THOMSON, J. W. 1984. American Arctic Lichens. The Macrolichens. Columbia University Press, NY.
———. 1997. American Arctic Lichens. The Microlichens. University of Wisconsin Press, Madison.
TUHKANEN, S. 1984. A circumboreal system of climatephytogeographical regions. Acta Botanica Fennica
127: 1–50.
USDA, NRCS. 2001. The PLANTS database, Version 3.1
(http://plants.usda.gov). National Plant Data Center,
Baton Rouge, LA.
WESTOFF, V. & E. VAN DER MAAREL. 1973. The BraunBlanquet approach, pp. 617–626. In R. H. Whittaker
(ed.), Ordination and Classification of Communities.
Junk, The Hague.
WINSLOW, M. A. 1992. Modeling paleoshorelines in geologically active regions: applications to the Shumagin
Islands, southwest Alaska, pp. 151–169. In L. L. Johnson (ed.), Paleoshorelines and Prehistory: An Investigation of Method. CRC Press, Ann Arbor, MI.
ms. received July 10, 2001; accepted Oct. 12, 2001.