Page  31 ï~~2002 THE MICHIGAN BOTANIST 31 BRYOPHYTES OF THE PICTURED ROCKS NATIONAL LAKESHORE, ALGER COUNTY, MICHIGAN, USA Janice M. Glime Department of Biological SciencesMichigan Technological University Houghton, MI 49931 ABSTRACT Field sampling for the present study includes relev6s of shoreline, coniferous forest, hardwood forest, mixed hardwood/coniferous forest, wetlands, and disturbed areas within the Pictured Rocks National Shoreline. This study adds 90 taxa of mosses, bringing the published total to 138 taxa (species and varieties) of mosses, and 18 liverworts, bringing that total to 38. The representative bryophytes in different habitats are discussed. Buxbaumia aphylla was found in the park for the first time, although it is relatively common in the Upper Peninsula of Michigan. Nine other uncommon and rare taxa were added to the list of known species in the park: Anomodon minor Calypogeja fissa (liverwort), Didymodon rigidulus, Herzogiella striatella, Hylocomium umbratum, Polytrichum pallidisetum, Rhabdoweisia crispata, Sphagnum wulfianum, and Tortula mucronifolia. INTRODUCTION The Pictured Rocks National Lakeshore occupies a location encompassing a variety of habitats, many of which are rare in Michigan. The Lakeshore lies in Alger County on the southern shore of Lake Superior, and therefore it occupies a significant portion of the undeveloped U.S. shoreline. Because of the effects of the lake, the northern position of the park in the Midwestern part of the country, and the alkaline sandstones, the park offers unique opportunities for finding plants that may occur nowhere else in the U.S., or at least nowhere else in the Midwest. Therefore, the preservation of selected habitats within the park, where such unusual flora and fauna can be expected, is important to meet part of the mission of the National Parks. Two French explorers, Radisson and Groseilliers, in the early 1660s, were probably the first Europeans to explore the Pictured Rocks area (Thwaites 1888). The first botanical description is attributed to Doty, based on his participation in the 1820 Cass Expedition (Doty 1895). But he stated only that "birch, beech and maple" grow along the top of the cliff. Subsequently, a number of botanical observations and collections of vascular plants have been reported (Dodge 1918; Fernald 1935; Cain 1962; Frederick, et al. 1977; Bach 1978; Crispin, et al. 1984). Read (1975) studied the vascular plants of Pictured Rocks, described the major vegetation types, and gave a detailed species checklist. Although there have been considerable investigations of vascular plant flora in the national parks nationwide, their bryophytes are still little known, particu

Page  32 ï~~32 THE MICHIGAN BOTANIST Vol. 41 larly in Midwestern national parks, being studied primarily in the western parks (Standley 1920; Porter 1932; Haring 1941, 1946; Persson & Weber 1958; Hong 1968, 1980; Hermann 1969, 1973, 1987; Magill 1976; Hoe 1979; Seyer 1979; Smith 1981; Showers 1982; Spence 1985; Blaney & Norris 1987). Few studies of bryophytes have addressed any park in the eastern half of the United States (e.g. Prior 1959; Ireland 1961). Only Isle Royale has been studied in any detail (Cooper 1912, 1913; Conklin 1914; Thorpe & Povah 1935; Hermann 1962; Rutkowski 1984; Meston 1985). In temperate and boreal regions bryophytes can play a very important ecological role in the forest ecosystem (Slack 1977, 1983; Glime 2001). Bryophyte communities are generally delimited by vascular plant communities, although bryophytes can have broader niches and form fewer communities within a habitat gradient than do vascular plants (Glime et al. 1982). The succession of the forest and the degradation of the environment will substantially influence the composition of the bryophyte flora in many different ways. Establishment of national parks provides the opportunity of not only protecting the natural habitat and forest but also conserving the bryophyte flora associated with the natural forest. Several bryologists have found particularly interesting bryophytes at Pictured Rocks National Lakeshore (Steere 1934; Conard 1938; Schuster 1949), but no comprehensive list of the taxa has ever been published. The interesting bryophytes already known from nearby areas, such as the rare Tetradontium brownianum, Cirriphyllum piliferum, and Timmia megapolitana at Tannery Falls in Alger County (Bowers 1987), likewise suggest that the park should harbor interesting species. The purpose of this survey was to assess the bryophyte vegetation from representative habitats to locate any interesting, endangered, or rare species, and to provide a checklist of the bryophytes known from the park. DESCRIPTION OF STUDY AREA The Pictured Rocks National Lakeshore occupies 71,397 acres of land along the southern shore of Lake Superior in the north-central section of the Upper Peninsula of Michigan, 460N latitude, 860W longitude. The most spectacular feature of the lakeshore is a 20 km shoreline of multicolored sandstone cliffs rising, at places, almost 60 m above the lake. Its sand dunes, reaching 90 m, are unique in the Upper Peninsula and provide unusual ridge and valley habitats supporting a variety of plant communities. Read (1975) described how the vegetation along the Lake Superior shoreline, the most attractive feature of the National Lakeshore, is strongly influenced by the moderating effects of the lake. The actual lake frontage may be divided into two types: beach strand-dune community and the sandstone cliff community. The beach strand-dune community is well developed on Sand Point, Miners Beach, Chapel Beach, and Twelve Mile Beach, the latter bordering our sampling corridor. We concentrated our sampling in a forested corridor 0.7 km wide and 10 km long, which borders the shoreline (Fig. 1), on 25-28 May 1991. The corridor in

Page  33 ï~~2002 THE MICHIGAN BOTANIST 33 AU SABLE.;:POINT LAKE GRAND SUPERIOR MARAIS TE HL KL GRA NDBL: Beaver Lake S AND-GP: Beaver Lake gravel pit HL: Hyde Lake KL: Kingston Lake Campground interior boundary. LB: Beaver Lake LBC: Little Beaver Lake Campground N LBL: Little Beaver Lake 0 3 LL: Legion Lake MB: Miners Beach Falls SMC: Sevenmile Creek exterior boundary SML: Sevenmile Lake TBE: Twelvemile Beach Campground east MUNISING TBW: Twelvemile Beach Campground west FIGURE 1. Map of Pictured Rocks National Lakeshore, Alger County, Michigan, showing locations of collection areas. For detailed locations, see Table 1. (Redrawn from Read 1975.) cludes coniferous forest, northern hardwood, hardwood with small coniferous component, hardwood with significant coniferous component, and pine barrens (NPS Assessment of Alternatives for Pictured Rocks National Lakeshore 1980). Although a variety of habitats exists in the park, the corridor is limited to forested areas, with only a few small woodland fen pools surrounded by Sphagnum, a gravel pit, and open trails and parking lots offering habitat variability. In addition to the corridor, collections were made at the cliff community of Miners Beach, since that community is the one most likely to include interesting and rare taxa, and earlier (1982) at Little Beaver Lake and Little Beaver Lake Campground. Additional records are from the literature. METHODS AND PROCEDURES Seven members of the Michigan Technological University Bryology Lab sampled by relev6s in the major habitats of the corridor and other areas, with more intense examination at Miners Beach. During the investigation, each taxon identifiable as potentially different in each vegetation type and habitat was collected and labelled. There were 1,500 specimens collected. Literature citations of species previously found in the park are indicated as such in Table 1. Scientific names of mosses follow the latest List of Mosses of North America North of Mexico provided by Anderson et al. (1990); the names for liverworts follow Stotler & Crandall-Stotler (1977) and Crum (1991). Nomenclature for vascular plants follows Gleason & Cronquist (1991). Voucher specimens of all bryophytes are in the Michigan Technological University Cryptogamic Herbarium (MCTC). RESULTS AND DISCUSSION To the list of known taxa, we have added 90 moss taxa and 18 liverwort taxa, bringing the totals to 136 and 38, respectively (Table 1). These numbers compare

Page  34 ï~~34 THE MICHIGAN BOTANIST Vol. 41 TABLE 1. Bryophyte species checklist of Pictured Rocks National Lakeshore. Species known from the literature are listed with reference in parentheses; nomenclature has been changed to conform with Anderson et al. 1990. Collections examined in this and the 1982 study have the following codes to denote collecting sites: BL: about 2 km southeast of Beaver Lake, west of the parking area about 1.5 km (460 33' N, 860 19' W, T48N R16W, Sec. 21) GP: road from H58 to the parking area about 3 km southeast of Beaver Lake, gravel pit north of road (460 32' N, 860 19' W, T48N R16W, Sec. 21) HL: about 1/2 km southeast of Hyde Lake (460 34' N, 860 16' W, T48N R16W, Sec. 14) KL: trail from Twelvemile Beach to Kingston Lake Campground (460 37' N, 860 15' W, T49N R15W, Secs. 24,25) LB: about 3 km southeast of Beaver Lake; parking area and along trail to Little Beaver Campground (460 33' N, 860 18' W, T48N R16W, Secs. 15, 16) LBC: Little Beaver Lake Campground, 1982 collections of Janice M. Glime (460 30' N, 860 30' W, T48N R17W, Sec. 18) LBL: Near Little Beaver Lake, 1982 collections of Janice M. Glime (470 30' N, 860 30' W, R48N R17W) LL: about 1/2 km E of Legion Lake, westernmost end of corridor (460 32' N, 860 21' W, T48N R16W, Sec. 30) MB: Miners Beach Falls (460 30' N, 860 32' W, T47N R18W, Secs. 2, 3) SMC: about 1 km south of the mouth of Sevenmile Creek, near NPS boundary markers on trees (460 37' N, 860 15' W, T49N R16W, Sec. 25) SML: about 1 km north northeast of Sevenmile Lake (460 35' N, 860 17' W, T48N R16W, Sec. 2) TBE: near the road between Twelvemile Beach Campground and the parking area ca 1 km to east (460 38' N, 860 13' W, T49N R15W, Sec 17) TBW: road about 1 km south of the parking area just east of Twelvemile Beach Campground, west side of the road (460 38' N, 860 13' W, T49N R15W, Secs. 17, 20) MOSSES Amblystegium varium (Hedw.) Lindb. (Miners Castle, Richards 1952) Anomobryum filiforme (Dicks.) Solms in Rabenh. (Conard 1938) Anomodon attenuatus (Hedw.) Huib. [JG13210] BL, KL, LB, LBL Anomodon minor (Hedw.) Fuirnr. [NL1585] BL, SMC Atrichum altecristatum (Ren. & Card.) Smyth & Smyth [JG13318] BL, HL, LB, SMC, SML Atrichum angustatum (Brid.) Bruch & Schimp. in B.S.G. GP, HL, KL, LBL, LL Aulacomnium palustre (Hedw.) Schwaegr. [JG13386a] KL Barbula convoluta var. convoluta Hedw. [JZ1172] MB, SML, TBE Barbula unguiculata Hedw. [JZ1068] BL, GP, TBE Bartramia pomiformis Hedw. [JG5496] LBC Brachythecium oxycladon (Brid.) Jaeg. [JZ1096] GP, HL, KL, LB, LL, SMC, SML, TBE Brachythecium plumosum (Hedw.) Schimp. in B.S.G. [JZ1063] SMC, TBE Brachythecium reflexum (Starke ex Web. & Mohr.) BSG (Grand Sable Dunes, Bowers 1987) [JG13216] BL, HL, LB, LBC, LBL, LL, MB, SML, TBE Brachythecium rivulare Schimp. in B.S.G. [JZ1162B] LBC, MB, SML Brachythecium salebrosum (Web. & Mohr) Schimp. in B.S.G. var. salebrosum (Grand Sable Dunes, Bach 1978) [JZ1195A] TBE, MB, SML, BL, KL, LB Brachythecium velutinum (Hedw.) Schimp. in B.S.G. var. velutinum [JZ1110C] KL, LB, SMC, SML Brotherella recurvans (Michx.) Fleisch. [JZ1077] HL, KL, LBC, LBL, SMC, SML, TBE Bryhnia novae-angliae (Sull. & Lesq. in Sull.) Grout (Grand Sable Dunes, Bowers 1987) [NL1528] MB Bryoerythrophyllum recurvirostre (Hedw.) Chen [JZ1171] MB Bryum argenteum Hedw. [JG13201] BL, LB, MB Bryum caespiticium Hedw. (Miners Castle, Richards 1952) [JG13432] MB Bryum capillare Hedw. [JZ1195C] SML Bryum lisae var. cuspidatum (Bruch & Schimp. in B.S.G.) Marg. [JZ1002] LB, MB Bryum pallens (Brid.) Sw. in Rohl. (rock crevices along shore, Nichols 1933) Bryum pseudotriquetrum (Hedw.) Gaertn. et al. [NP224] GP, LBL, MB, SML

Page  35 ï~~2002 THE MICHIGAN BOTANIST 35 TABLE 1. (Continued) Buxbaumia aphylla Hedw. [NP197] KL, SMC Callicladium haldanianum (Grev.) Crum [JG13317] BL, GP, HL, KL, LB, LBL, SMC, SML, TBE, TBW Campylium hispidulum (Brid.) Mitt. [NP2] LB, SMC Campylium polygamum (Schimp. in B.S.G.) C. Jens. [NP19] LB Catoscopium nigritum (Lake Superior shore, Nichols 1933; Conard 1938) Ceratodon purpureus (Hedw.) Brid. [JG13269] BL, GP, HL, KL, LB, MB, SML, SMC, TBE Climacium dendroides (Hedw.) Web. & Mohr [NP223] LBC, MB Cratoneuron filicinum (Hedw.) Spruce [JZ1167A] MB Desmatodon obtusifolius (Schwaegr.) Schimp. (sandstone cliffs, Steere 1934; Conard 1938; Miners Castle, Richards 1952) [JZ1178] MB Dichodontium pellucidum (Hedw.) Schimp. (base of lake bluffs, Nichols 1933; Conard 1938) [JZ1167B] KL, MB Dicranella cerviculata (Hedw.) Schimp. (moist sandstone, ledge of cliff, Miners Castle, Crum & Miller 1968) Dicranella grevilleana (Brid.) Schimp. (moist gravelly slopes at Miners Falls, Nichols 1933) Dicranella heteromalla (Hedw.) Schimp. (Grand Sable Dunes, Bach 1978) [JZ1220] BL, HL, LB, LBC, LL, SMC Dicranumflagellare Hedw. [JG13281] KL, SML, TBE, TBW Dicranumfuscescens Turn. [JZ1039B] HL, KL, SML, TBE, TBW Dicranum montanum Hedw. [JZ1060] BL, GP, HL, KL, LB, LBL, LL, MB, SMC, SML, TBE, TBW Dicranum polysetum Sw. (Grand Sable Dunes, Bowers 1987) [JG13407] GP, KL, LBC, TBE, TBW Dicranum scoparium Hedw. (Nichols 1933) [JZ1037] BL, GP, HL, KL, LB, SMC, SML, TBE, TBW Dicranum undulatum Brid. [JG13414] KL Dicranum viride (Sull. & Lesq. in Sull.) Lindb. [JZ1027] BL, HL, KL, LB, SMC, SML, TBE Didymodon rigidulus var. rigidulus Hedw. [JG13446] MB Diphysciumfoliosum (Hedw.) Mohr [NP33] LB Ditrichum flexicaule (Schwaegr.) Hampe (Grand Sable Dunes, Bach 1978) Encalypta ciliata Hedw. (Miners Falls, Steere 1934) [NL1718] MB Eurhynchium pulchellum (Hedw.) Jenn. [JM1162] LBC, TBE Fissidens dubius P. Beauv. [NP51B] BL Fissidens osmundioides Hedw. LBL Fontinalis hypnoides var. duriaei (Schimp.) Husn. [JG5536] LBL Funaria hygrometrica Hedw. [JG13420] MB Gymnostomum aeruginosum Sm. [JZ1170B] MB Herzogiella striatella (Brid.) Iwats. [JG13255] TBE Hygroamblystegium fluviatile (Hedw.) Loeske [JG13434] MB Hygroamblystegium noterophilum (Sull. & Lesq. in Sull.) Warnst. (Miners Falls, Steere 1934) Hygroamblystegium tenax (Hedw.) Jenn. (Miners Falls, Nichols 1935) [JG13423] KL, LB, MB Hygrohypnum luridum (Hedw.) Jenn. (along lakeshore & Miners Falls, Nichols 1933) [JZ1175] MB, SML Hylocomium splendens (Hedw.) Schimp. in B.S.G. [NL1668] KL, LBL, TBE Hylocomium umbratum (Hedw.) Fleisch. in Broth. [JG5533] LBL Hymenostylium recurvirostre (Hedw.) Dix. (Miners Castle, Nichols 1933; Richards 1952) [JZ1174A] KL, MB, TBE Hypnum cupressiforme L. (lake shore, Nichols 1933) Hypnum imponens Hedw. [JG13379] KL, SMC, TBE Hypnum pallescens (Hedw.) P. Beauv. var. pallescens [JG13330] BL, GP, KL, LB, LBL, SMC, SML, TBE, TBW Leptobryum pyriforme (Hedw.) Wils. [JZ1174C] MB, SMC Leptodictyum humile (P. Beauv.) Ochyra [NL1562] LB Leptodictyum riparium (Hedw.) Warnst. [JG5524] LBL Leskeella nervosa (Brid.) Loeske [JG13215] BL, GP, HL, LB, TBE, SML Leucobryum glaucum (Hedw.) Angstr. in Fries [JG13285] KL, TBE Leucodon brachypus var. andrewsianus Crum & Anderson (Miners Rock, Bowers 1987) [NL1556] BL, LB (Continued)

Page  36 ï~~36 THE MICHIGAN BOTANIST Vol. 41 TABLE 1. (Continued) Meesia triquetra (Richt.) Angstr. (rock crevices near lake shore, Nichols 1933) Mielichhoferia mielichhoferiana (Funck in Hook.) Loeske var. mielichhoferiana (sandstone cliffs, Nichols 1933; Conard 1938; near Miners Castle, Crum & Miller 1968; Miners Rock, Crum & Anderson 1981) Mnium spinulosum Bruch & Schimp. in B.S.G. [JZ1198] SMC, SML Mnium thomsonii Schimp. [JZ1190] LL, SML Myurellajulacea (Schwaegr.) Schimp. in B.S.G. (Lake Superior shore, Nichols 1933; Miners Castle, Richards 1952) Myurella sibirica (C. Muill.) Reim. (Miners Castle, Richards 1952) Neckera pennata Hedw. [JZ1029A] BL, KL, SMC Orthotrichum sordidum Sull. & Lesq. in Aust. [JZ1025B] BL, GP, KL, LBC, SMC, TBW Orthotrichum speciosum var. elegans (Schwaegr. ex Hook. & Grev.) Warnst. (Miners Rock, Bowers 1987) Paraleucobryum longifolium (Hedw.) Loeske [NP53A] BL Philonotisfontana (Hedw.) Brid. var. fontana (Miners Castle, Richards 1952) [JG13457] MB Plagiomnium ciliare (C. M.) T. Kop. [JG13223] BL, KL, LB, LBL, LL Plagiomnium cuspidatum (Hedw.) T. Kop. [JZ1106B] BL, GP, LB, LL, MB, SMC, TBE, TBW Plagiomnium ellipticum (Brid.) T. Kop. [JZ1008] LB Plagiothecium cavifolium (Brid.) Iwats. [JG13317a] HL, LB, LL, SMC, SML, TBW Plagiothecium laetum Schimp. in B.S.G. [NP176] BL, GP, HL, KL, LB, LL, MB, SML, TBE?Platydictya subtilis (Hedw.) Crum [JZ1189A] SML Platygyrium repens (Brid.) Schimp. in B.S.G. (Grand Sable Dunes, Bowers 1987) [JZ1188] BL, HL, LB, MB, SMC, SML, TBE, TBW Platyhypnidium riparioides (Hedw.) Dix. (Miners Falls, Steere 1934) Pleurozium schreberi (Brid.) Mitt. [JG13250] GP, KL, LB, LBL, SMC, TBE, TBW Pogonatum dentatum (Brid.) Brid. (gravelly lake bluffs, Nichols 1933; Conard 1938) Pohlia annotina (Hedw.) Lindb. (Miners Castle, Crum & Miller 1967) Pohlia elongata Hedw. var. elongata (Munising Falls, Miners Falls, Steere 1934) Pohlia nutans (Hedw.) Lindb. [JG13259] HL, KL, SMC, SML, TBE Pohlia proligera (Kindb. ex Breidl.) Lindb. ex Arnell (Crum & Miller 1967) Pohlia wahlenbergii (Web. & Mohr) Andrews (crevices of cliffs, Nichols 1933) [JG1170A] MB Polytrichastrum alpinum (Hedw.) G. L. Sm. var. alpinum (woods along lake bluffs, Nichols 1933; Conard 1938) [JM1174] HL, TBE, TBW Polytrichum commune Hedw. var. commune [JZ1021] LB, SML Polytrichum juniperinum Hedw. (Grand Sable Dunes, Bach 1978) [JG13305] LB, MB, SMC, TBE, TBW Polytrichum longisetum Brid. [JZ1221A] LL, SML Polytrichum ohioense Ren. & Card. [JZ1203A] HL, KL, LBL, SMC, SML, TBE Polytrichum pallidisetum Funck [JZ1119A] GP, HL, LL, SMC, SML Polytrichum piliferum Hedw. [JG13248] GP, KL, SMC, TBE Pseudobryum cinclidioides (Hub.) T. Kop. (Grand Sable Dunes, Bowers 1987) Pseudoleskea radicosa (Mitt.) Mac. & Kindb. (Grand Sable Dunes, Bowers 1987) Ptilium crista-castrensis (Hedw.) De Not. [JG13251] KL, TBE, TBW, SMC Pylaisiella selwynii (Kindb.) Crum et al. [JG13211] BL, KL, LB, LBC, SMC, TBE, TBW Racomitrium canescens (Hedw.) Brid. var. canescens (Grand Sable Dunes, Bach 1978) [NP265] GP Rhabdoweisia crispata (With.) Lindb. [JG13426] KL, MB Rhizomnium pseudopunctatum (Brunch & Schimp.) T. Kop. (Grand Sable Dunes, Bowers 1968) Rhizomnium punctatum (Hedw.) T. Kop. [NP222] MB, SMC, SML Rhytidiadelphus triquetrus (Hedw.) Warnst. (Grand Sable Dunes, Bach 1978) [JZ1165] LB, LBC, MB, TBE Saelania glaucescens (Hedw.) Broth. in Bomanss. & Broth. [JZ1174B] KL, MB Sanionia uncinata (Hedw.) Loeske var. uncinata [JZ1090] KL, LB, MB, SMC, TBE, TBW Schistidium apocarpum (Hedw.) Bruch & Schimp. in B.S.G. (Miners Castle, Richards 1952) Scopelophila ligulata (Spruce) Spruce (sandstone outcrop, Miners Castle, Crum & Miller 1968) Seligeria donniana (Sm.) C. Mull. (Miners Falls, Steere 1934) Sphagnum angustifolium (C. Jens. ex Russ.) C. Jens. in Tolf [JG13415] KL Sphagnum capillifolium (Ehrh.) Hedw. [JG13346] KL, SMC

Page  37 ï~~2002 THE MICHIGAN BOTANIST 37 TABLE 1. (Continued) Sphagnum centrale C. Jens. in Arnell & C. Jens. [JG13385] KL Sphagnum cuspidatum Ehrh. ex Hoffm. [JG13401] KL Sphagnum fallax (Klinggr.) Klinggr. [JG13402] KL Sphagnum fimbriatum Wils. in Wils. & Hook. f. in Hook. f. (Conard 1938) Sphagnum girgensohnii Russ. [JG13398] KL Sphagnum papillosum Lindb. [JG13411] KL Sphagnum rubellum Wils. [JG13413] KL Sphagnum squarrosum Crome [JG13388] KL, LBC Sphagnum wulfianum Girg. [JG13400] KL Steerecleus serrulatus (Hedw.) Robins. [JG13245] BL Tetraphis pellucida Hedw. [JG13368] BL, KL, LB, LBC, SMC, SML, TBE, TBW Thuidium delicatulum (Hedw.) Schimp. in B.S.G. [NL1721] MB Thuidium recognitum (Hedw.) Lindb. [JG13278] LB, MB, TBE Tortula mucronifolia Schwaegr. [JG13441] MB, TBW Ulota crispa (Hedw.) Brid. [NL1763] GP, HL, KL, LBC, SMC LIVERWORTS Anastrophyllum michauxii (Web.) Buch et Evans (gravelly lake bluff & cliff crevices, Nichols 1933) Anthelia juratzkana (Limpr.) Trev. (Conard 1938; sandstone cliff, east of Miners Castle, Crum & Miller 1968) Bazzania trilobata (L.) S. Gray [JG13351] HL, KL, LBC, LBL, SMC, TBE, TBW Blasia pusilla L. (gravelly lake bluffs, Nichols 1933; Conard 1938) Blepharostoma trichophyllum (L.) Dum. [JG5497] LBC Calypogejafissa (L.) Raddi [JG13230] LB, LBC Calypogeja integristipula Steph. [JG5503] LBC Calypogeja muelleriana (Schiffn.) K. Mull. [JG13279] TBE Cephalozia bicuspidata (L.) Dum. (gravelly lake bluff, Nichols 1933) Cephalozia lunulifolia (Dum.) Dum. [JG5519] LBL Conocephalum conicum (L.) Lindb. (Miners Falls, Steere 1934) [JG5517] LBL Frullania bolanderi Aust. (Conard 1938) [JG13222] GP, KL, LB, LBL, SMC, SML Frullania eboracensis Gott. [JZ1097B] SMC Jamesoniella autumnalis (DC.) Steph. [NP228] GP, KL, SMC, SML, TBE, TBW Jungermannia exsertifolia subsp. cordifolia (Dum.) Vana (lake bluffs, Nichols 1933; Conard 1938) Jungermannia hyalina Lyell (rock ledges along shore, Nichols 1933; Steere 1934) Jungermannia leiantha Grolle (Conard 1938) Jungermannia sphaerocarpa Hook. (sandstone cliff, Miners Castle, Crum & Miller 1968) Lepidozia reptans (L.) Dum. [JZ1033A] BL, KL, LBC, TBE, TBW Lophocolea heterophylla (Schrad.) Dum. [JZ1046] KL, LB, LBL, SML, TBE Lophocolea minor Nees [JG13444] MB Lophozia alpestris (Schleich. ex Web.) Evans (gravelly lake bluffs, Nichols 1933) Lophozia attenuata (Mart.) Dumort. [JZ1082] KL, TBE, TBW Lophozia badensis (Gott. ex Rabenh.) Schiffn. (gravelly lake bluffs, Nichols 1933; Conard 1938) Lophozia gillmanii (Aust.) Schust. (gravelly lake bluffs, Nichols 1933) Lophozia ventricosa (Dicks.) Dumort. (base of cliffs, Nichols 1933) Pellia epiphylla (L.) Corda (gravelly lake bluffs, Nichols 1933) Plagiochila porelloides (Torrey ex Nees) Lindenb. [JG5515] LBL Porella platyphylla (L.) Pfeiff. [JG13234] BL, LB, SML Preissia quadrata (Scop.) Nees (Miners Castle, Richards 1952) [JZ1164] KL, MB Ptilidium ciliare (L.) Hampe [JG13301] HL, KL, TBE, TBW Ptilidium pulcherrimum (G. Web.) Hampe [JG13366] BL, KL, LB, LBL, SML, TBE, TBW Radula complanata (L.) Dum. [JG1323] BL, LB, SML Scapania irrigua (Nees) Gott. et al. (rock ledges, Nichols 1933) Scapania nemorosa (L.) Dum. (Steere 1934) Scapania saxicola Schust. (sandstone cliff, east of Miners Castle, Crum & Miller 1968) Trichocolea tomentella (Ehrh.) Dum. [JG5487] LBC Tritomaria exsecta (Schrad.) Loeske (gravelly lake bluff, Nichols 1933)

Page  38 ï~~38 THE MICHIGAN BOTANIST Vol. 41 with over 400 mosses and liverworts reported for the state of Michigan. The bryophyte flora is described below based on the vegetation types according to the 1980 NPS Assessment of Alternatives for Pictured Rocks National Lakeshore, with locations noted on Figure 1. The corridor represents only a small percentage of the likely taxa in the park because it is mostly forested and lacks the habitats that are rich in species. The alkaline wetlands may harbor taxa that occur nowhere else in the Upper Peninsula, and because of their northern location may harbor taxa that are unknown elsewhere in the state. Furthermore, the most interesting and unusual habitats are the alkaline sandstones along the rocky shore, and these have barely been explored by any bryologists. A boat trip along the cliffs could reveal some very interesting finds. Lake shoreline Bryophytes of the lake shoreline grow in a unique habitat affected by both the lake and the forest. Relatively more diverse microhabitats are found in this area. Near the road between Twelvemile Beach Campground and the parking area about 1 km to the east (TBE) there are three types of habitats: foredune, valley between dunes, and second dune. The dune face is a very unstable habitat and the only species found there abundantly is Polytrichum piliferum (Fig. 2). This species is a widely distributed moss and most commonly grows in disturbed, exposed, and dry habitats. In the Netherlands, it is an important component on inland dunes (Plius 1994) and on the shores of Lake Superior it plays an important role in dune stabilization (Marsh & Koerner 1972). Bach (1978) also found Brachythecium salebrosum, Dicranella heteromalla, Ditrichum flexicaule, Racomitrium canescens, Rhytidiadelphus triquetris, and Polytrichum juniperinum on the park dunes, but did not report any Polytrichum piliferum. Mosses such as these are able to trap blowing sand, thus helping to stabilize the dunes (Schulten 1985) At the top of the dune there is an overstory including Pinus resinosa, Betula papyrifera, Acer saccharum, and other hardwood and coniferous trees. On the forest floor the common bryophytes Brachythecium oxycladon, B. velutinum var. velutinum, Callicladium haldanianum, Dicranum scoparium, D. montanum, D. polysetum, Ptilium crista-castrensis, Sanionia uncinata, and Pleurozium schreberi are frequently found; on tree bases Brachythecium reflexum, Dicranum viride, and Hypnum pallescens; and on rotten logs Brotherella recurvans, Lophocolea heterophylla (liverwort), and Ptilidium pulcherrimum (liverwort). The valley between the foredune and second dune is a habitat with more shade and taller trees. The overstory is dominated by Acer saccharum, Betula papyrifera, and occasionally Pinus resinosa. The understory is mostly Pinus strobus saplings. The forest floor is covered by such vascular plants as Gaultheria procumbens, Maianthemum canadense, Lycopodium clavatum, Pteridium aquilinum, Trientalis borealis, and Vaccinium sp. Bryophytes are more abundant in the protective microhabitat of the valley than on the foredune. One of the noticeable habitat differences from that of the foredune is that there are more rotten logs and stumps and a thicker litter layer in the valley. Among the bryophytes are those growing on rotten logs and stumps: Callicladium haldanianum, Bazza

Page  39 ï~~2002 THE MICHIGAN BOTANIST 39 2002 THE MICHIGAN BOTANIST 39 FIGURE 2. Shoreline along Twelvemile Beach, Pictured Rocks National Lakeshore, Alger County, Michigan, showing Polytrichum piliferum creating small hummocks (see arrow) that stabilize the beach. Photograph by the author, late May, 1991. nia trilobata (liverwort), Brotherella recurvans, Lophocolea heterophylla (liverwort), Lepidozia reptans (liverwort), Plagiothecium laetum, Sanionia uncinata, and Tetraphis pellucida; species found on soil: Dicranum scoparium, Hypnum pallescens, Ptilium crista-castrensis, Platygyrium repens, Thuidium recognitum, and Barbilophozia attenuata (liverwort). On sandy soil at the top of the second dune, in addition to the species found on the foredune, are Brachythecium salebrosum, Hylocomium splendens, Hypnum imponens, Plagiomnium cuspidatum, Plagiothecium laetum, and Rhytidiadelphus triquetrus. Coniferous forest A Jack pine forest (Pinus banksiana) was sampled about 1 km south of the parking area just east of Twelvemile Beach Campground (TBW). This is an open and dry habitat with a few Acer saccharum saplings in the understory. The ground is covered sparsely by the lichens Cladina spp. and Cladonia spp., and vascular plants Epigaea repens, Maianthemum canadense, Pteridium aquilinum, and Vaccinium sp. The habitat is uniform, with few bryophyte taxa, the representative ones being Dicranum flagellare, D. fuscescens, D. montanum, D. polysetum, D. scoparium, Plagiothecium cavifolium, Pleurozium schreberi, Polytrichum juniperinum, and Ptilium crista-castrensis.

Page  40 ï~~40 THE MICHIGAN BOTANIST Vol. 41 Hardwood forests The hardwood forest, associated with few coniferous trees, was sampled at several locations: LB, BL, HL, LL, SML (See Table 1). The habitat is more shady and moist than the coniferous forest, with a dominant overstory of Acer saccharum, Fagus grandifolia, and Acer pensylvanicum; understory saplings include Acer saccharum, Fagus grandifolia, and Acer pensylvanicum; the sparse ground cover includes vascular plants Actaea sp., Aralia nudicaulis, Claytonia caroliniana, Dicentra spp., Carex spp., Clintonia borealis, Dryopteris intermedia, Erythronium americanum, Lycopodium (sensu lato) spp., Maianthemum canadense, Milium effusum, Mitchella repens, Osmorhiza spp., Oxalis sp., Sambucus pubens, Smilacina racemosa, Trientalis borealis, and Viola sp. There are more species of bryophytes in the hardwood forest than in the coniferous forest. Forest floor taxa include Amblystegium varium, Bazzania trilobata (liverwort), Brachythecium sp., B. reflexum, Callicladium haldanianum, Dicranella heteromalla, Dicranum scoparium, Hypnum pallescens, Mnium thomsonii, Plagiothecium spp., and Jamesoniella autumnalis (liverwort); on tree bases and tree bark, Brachythecium reflexum, Frullania bolanderi (liverwort), Hypnum pallescens, Leskeella nervosa, Orthotrichum sp., Platygyrium repens, Porella platyphylla (liverwort), Ptilidium pulcherrimum (liverwort), and Pylaisiella selwynii. Mixed hardwood and coniferous forests The mixed hardwood and coniferous forest is a widely distributed major vegetation type in the park. We sampled two major locations: SMC and KL (See Table 1). The overstory species are represented by Abies balsamea, Acer saccharum, Betula papyrifera, Fagus grandifolia, and Tsuga canadensis; understory by saplings of Acer pensylvanicum and A. saccharum; and ground cover by Aralia nudicaulis, Coptis trifolia, Dryopteris intermedia, Lycopodium spp., Maianthemum canadense, Oxalis sp., and Trientalis borealis. The habitat characters and bryophyte species of this type of forest are not very different from those of the hardwood forest, with common forest floor taxa including Brachythecium sp., B. reflexum, Callicladium haldanianum, Ceratodon purpureus, Climacium dendroides, Dicranum flagellare, D. montanum, D. scoparium, D. viride, Hypnum pallescens, Plagiomnium cuspidatum, Plagiothecium spp., Pleurozium schreberi, Pohlia spp., Tetraphis pellucida, and the liverworts Bazzania trilobata, Jamesoniella autumnalis, Lepidozia reptans, and Lophocolea heterophylla; Brotherella recurvans, Frullania bolanderi (liverwort), Neckera pennata, Orthotrichum speciosum, Platygyrium repens, and Ptilidium pulcherrimum (liverwort) occur on bark. Wetlands Wetlands have been particularly important for special concern, threatened, and endangered vascular plants in the park (Crispin et al. 1984). We sampled several wetlands west of the trail from Twelvemile Beach to Kingston Lake

Page  41 ï~~2002 THE MICHIGAN BOTANIST 41 Campground (KL) where the overstory is dominated by Betula alleghaniensis, Larix laricina, Picea mariana, and Tsuga canadensis, and the groundcover by Andromeda glaucophylla, Chamaedaphne calyculata, Eriophorum sp., Kalmia polifolia, Ledum groenlandicum, and Vaccinium sp. (cranberry). Ten species of Sphagnum were identified: S. angustifolium, S. capillifolium, S. centrale, S. cuspidatum, S. fallax, S. girgensohnii, S. papillosum, S. rubellum, S. squarrosum, and S. wulfianum. Near parking lots and trail entrances The open, sandy habitat of the borders of parking lots and entrances to trails harbors a small group of light-tolerant, short mosses. These generally include Barbula spp., Bryum argenteum, Bryum spp., and Ceratodon purpureus. Rock cliffs at Miners Beach Falls Miners Beach Falls (MB) is on seepy rock cliffs next to Lake Superior. Because of its diverse microhabitats and topography it is the most interesting place to find many rare and unusual bryophytes that were scarcely found elsewhere in the park. Aquatic and wet habitat species included Brachythecium rivulare, Cratoneuron filicinum, Hygroamblystegium fluviatile, Hygrohypnum luridum, Lophocolea minor (liverwort), Philonotis fontana, Pohlia wahlenbergii, Preissia quadrata (liverwort), Rhizomnium pseudopunctatum (considered by Bowers, 1968, to be rare in the Upper Peninsula), and R. punctatum. Other interesting species found on sandstone here are Barbula convoluta, Bryoerythrophyllum recurvirostre, Desmatodon obtusifolius, Dichodontium pellucidum, Encalypta ciliata, Funaria hygrometrica, Gymnostomum aeruginosum, Hymenostylium recurvirostre, Leptobryum pyriforme, and Saelania glaucescens. Interesting and rare species Buxbaumia aphylla (Bug-on-a-stick) was found in the park for the first time, on the trail from Twelvemile Beach to Kingston Lake Campground on a steep, sandy southwest-facing hillside (KL) that is a deforested site with a few Populus trees scattered on the mostly unvegetated, sandy slope. Ground cover is mostly the lichens Cladina and Cladonia spp., and mosses Bryum sp., Ceratodon purpureus, Dicranum montanum, Pleurozium schreberi, Plagiomnium ciliare, and Polytrichum piliferum. Buxbaumia aphylla also occurs beside the trail near the beach south of the mouth of Sevenmile Creek. Buxbaumia aphylla has been found abundantly in several localities in Houghton, Keweenaw, and Baraga Counties in Upper Michigan, but has appeared on the working lists for protection in both Wisconsin (1993, unpub.) and the US Forest Service Globally Rare for USA and Canada (1996). It is a pioneer of disturbed, acid, sandy or clayey soil, often on the banks of roads or in woodlands, exposed or in partial shade in moist forest and also dry, open woods, often successional to fire. Another species that has a morphology somewhat similar to that of Buxbaumia is Diphyscium foliosum (grain of wheat moss). This dioecious moss was

Page  42 ï~~42 THE MICHIGAN BOTANIST Vol. 41 found only in a sample along the trail to Little Beaver Campground about 3 km southeast of Beaver Lake (LB). Although seemingly rare in the park, the species is common in the Upper Peninsula. Diphyscium usually grows on shaded soil or humus, especially on the banks of woodland trails. This time we only found the male gametophyte-a leafy plant with no capsule. To the uncommon species reported in the literature, we have added the mosses Anomodon minor, Didymodon rigidulus, Herzogiella striatella, Hylocomium umbratum, Polytrichum pallidisetum, Rhabdoweisia crispata, Sphagnum wulfianum, and Tortula mucronifolia, and the liverwort Calypogeja fissa (Wisconsin Natural Heritage Working List 1993, unpub.), totalling 9 species. From the literature (see Table 1), we can include Dicranella cerviculata (Wisconsin Natural Heritage Working List 1993, unpub.), D. grevilleana, Ditrichum flexicaule (Wisconsin Natural Heritage Working List 1993, unpub.), Hygroamblystegium noterophilum, Meesia triquetra (noted by Crum 1973 as extremely rare), Mielichhoferia mielichhoferiana (a rare copper moss; Michigan Natural Features Inventory compilation, 1982 unpub.; U. S. Forest Service Working Copy, 1996 unpub.), Pohlia annotina, Pogonatum dentatum, Pseudobryum cinclidioides, Pseudoleskea radicosa, Racomitrium canescens (the latter confirmed by our collections), Scopelophila ligulata (found only in this one collection in Michigan), and Seligeria donniana (Wisconsin Natural Heritage Working List 1993, unpub.) as rare or uncommon. To the liverworts, we can add from the literature Anastrophyllum michauxii, Anthelia juratzkana, Jungermannia exsertifolia subsp. cordifolia (Wisconsin Natural Heritage Working List 1993, unpub.), J. hyalina, J. sphaerocarpa, Lophozia alpestris, L. badensis, L. gillmanii, L. ventricosa, Scapania irrigua, and S. saxicola (known from one other collection in Michigan; Wisconsin Natural Heritage Working List 1993, unpub.). Most of these liverworts are not known from the Lower Peninsula. However, of these bryophytes only Mielichhoferia mielichhoferiana appears on the list of Rare Bryophytes of Michigan (Gereau & Crispin for Michigan Natural Features Inventory, 1982 unpub.). Herzogiella striatella is found on rotten birch logs associated with Hypnum pallescens, near the road between Twelvemile Beach Campground and the parking area about 1 km to the east. Being a northern species, it is unknown in the Lower Peninsula, but occurs at Tahquamenon Falls (Crum 1973). It is not common in the Upper Peninsula. Hylocomium umbratum, found in 1982 in the forest at Little Beaver Lake, is otherwise known in Michigan only from Tahquamenon Falls and one other site. Plagiomnium ciliare seems to be known in the Upper Peninsula only from Chippewa and Ontonagon Counties (Darlington 1964). Pseudoleskea radicosa is thus far known only from Keweenaw and Ontonagon Counties and Pictured Rocks in Michigan (Darlington 1964). Calypogeja fissa is previously known from only one locality in the Upper Peninsula, in the Keweenaw Peninsula (Glime & Slavick 1985). Other uncommon species found in the park are Ditrichum flexicaule at Grand Sable Dunes (Bach 1978), and Leptobryum pyriforme (BL). The species of Ditrichum and Leptobryum occur in disturbed habitats and therefore their habitat cannot easily be preserved; although not rare, they are not common in

Page  43 ï~~2002 THE MICHIGAN BOTANIST 43 the Upper Peninsula. Pictured Rocks (at about 1 km south of the mouth of Sevenmile Creek, near NPS boundary markers on trees; SMC) and the sand barrens of Schoolcraft County seem to be the only published locations for Polytrichum pallidisetum in Michigan, although Crum & Anderson (1981) reported it as common in the Great Lakes area. Racomitrium canescens grows on sandy soil; although it is very widespread in the northern hemisphere, it is uncommon in Michigan. We found it only along the road from road H58 to the parking area about 3 km southeast of Beaver Lake, gravel pit north of the road (GP). CONCLUDING REMARKS The bryophyte flora of the Pictured Rocks National Lakeshore is a protected habitat overlying alkaline sand. As a result, several species occur here that are difficult to find elsewhere. Most of these somewhat rare taxa are to be found on the lakeshore among the calcareous sandstone cliffs, but a few are found on the sandy forest soils, wetlands, and elsewhere. Another habitat with less common species is the bog/fen system. Among the bryophytes in this habitat, Sphagnum wulfianum is uncommon in the Upper Peninsula. Many wetlands exist in the park and remain to be explored for rare bryophytes. The lakeshore habitat, especially along the trail from Twelvemile Campground west, should be preserved. This beach area now has a beautiful hiking trail with numerous orchids, including a large, dense patch of Cypripedium acaule, among others, and the bryophytes Tortula mucronifolia and Buxbaumia aphylla. It is likely that many more bryophytes exist in the park but have never been identified. It would be beneficial from a phytogeographical perspective to do a thorough survey of the entire park to document those taxa that might be living in this unusual alkaline habitat in the transition between the boreal forest and the mixed deciduous forest typical further south. Pictured Rocks National Lakeshore represents a unique location not only in Michigan, but also in the United States, where one can find northern alkaline species. The Lake Superior Basin harbors many interesting species that seem to be disjunct from western communities, and surely many of these disjunct taxa remain to be discovered along the sandstone cliffs and in the wetland areas of Pictured Rocks National Lakeshore. ACKNOWLEDGMENTS The National Park Service, Pictured Rocks National Lakeshore, funded this project. Yenhung Li identified the species of Sphagnum. Jinkun Zhang, Janet Marr, and Nadine Plante helped with fieldwork and identifications. Howard Crum graciously verified many of the bryophytes, yet claimed to have "done nothing" when invited to be a co-author. YingPing Sheng, Na Li, and Chang-Liang Liao helped with the fieldwork.

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