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320 THE CANADIAN FIELD-NATURALIST Vol. 111

The alpicola Dix. New to Alberta and the Yukon Territory with a Discussion of its Range and Comments on Related Species

P. M. ECKEL

Clinton Herbarium, Buffalo Museum of Science, Buffalo, New York 14211-1293.

Eckel, r. 1997. The moss Tortella alpicola Dix, new to Alberta and the Yukon Territory with a discussion of its range and 1 . comments on related species. Canadian Field-Naturalist 111(2): 320--322. Tortclla alpicola Db.., once considered rare as only a few isolated stations had been recorded in North America, is shown to have a more widespread distribution. Characteristics distinguishing this species from the common and widespread Tortclla fragilis and T. IOrtuosa arc presented ina key. Key Words: Tortella alpicola, Tortella fra gllis, Tortella tortuosa, Tortclla nitlda, Alberta, Canada, Yukon Territory, rare bryophytcs, moss.

Among the collections of undetermined bryophyte specimen from Hawaii in the distribution under the material of the Geography Department of the name val. tortelloides (Greene) University o f Calgary (those not se nt to the Zand. & Hoe, as Zander and Hoe considered it only Provincial Museum of Alberta in Edmonton in 1995 marginally distinct from Tortella fragilis (Hook. & and examined subsequent to that year), specimens of Wils in Drumm.) Limpr. After study of material of Tortella alpicola Dix. came to light. The North the genus Tortella from throughout Arctic North American distribution of this species, due to taxo­ America, Zander (m anuscript in preparation) con­ nom ic difficulties, has not yet been adequately docu­ cluded that the variety was a good species, as origi­ mented. The University of Calgary specimens, and nally determined by Robinson. three others seen from Canada at other herbaria, rep­ Following Zander's decision, Eckel (1991) pub­ resent the present known distribution of the species lished the discovery of the species in the contermi­ in that country (herbarium code throughout follows nous United States from the sta te of Wyoming, that des ignated by Holmgren et al. 1990) : including reference to a station in the Northwest Territories of Canada. The taxon was discussed Tortella alpicola : again by Hyvoenen (1991), who indicated, based on Alberta: Red Deer River Watershed, provincial campsite at j unction of James River and Willson Creek on his examination of specimens of Tortella fragilis Coch rane -Nordegg Forestry Trunk Road. approx. from the southern hemisphere, that Tortella tortel­ 5J048'N, 115°03'W, elev. 4400 feet. June 23. 1961, C. loides should be plac ed under Tortella fragilis as a D. Bird 6249 ; with En calypta vulgaris (Geography variety again. My examination of specimens at H Dep artment, University of Cal gary): Bow River from Ti erra del Fuego collected by H. Roivainen, Watershed, 3 miles WSW of Exshaw, SWI7-24-9-W5, some of which were annotated by Hyvoenen, has 51°03'N, 115°13'W, elev. 4300 ft., Upper Foothills shown all of these specimens to be Tortella fragilis Forest, north-facing slope with steep limestone expo­ val. [ragili s. None of them were what is indicated sures, on May 19, 1976, Glennis Lewis no. 36, with here as Tortella alpicola. Other specimens at CHR, Encalypta rhaptocarpa and Gymnoslomumaerugino sum H, NY and ALTA of both Tortella [ragili s and T. (BUF); Jasper National Park, Dryas drummondii flat 2-3 tortuosa collected in the southern hemi sphere were miles below Athabaska Glacier, July 23, 1959, W. A. Weber B-2278 (COLO). all either correctly determined or were species other Northwest Territories: Ellesmere Island , lake cast of than T. alp icola. So far, the only known southern Taconite Inlet, 82°52'N, 78°05'W, cliff of metamorphic hemisphere stations for the latter species are in rock, 50 rn, May 21,1969, G. Brassard 4136 (CANM). Antarctica. The species has been found in Colombia Yukon Territory: Selwyn Mountains. Itsi Range. North end (Churchill and Linares 1995), and may prove to be a of unnamed lake , south of Fuller Lake , 62-57 'N, member of the Andean flora . Zander (1993) reported 130--09'W. 1600 m, in mesic Betula pumila-Vaccinium an earlier name for Tortella tortelloides based on the uliginosum dominated tundra on cast and north facing type of T. alpicola Dixon (1930), collected from a slopes, siliceous rock outcrops of granite and slate pre­ station in the Himalayas. sent, with small amounts of calcareous rock cropping As a result of my examination of numerous speci­ out, August 10, 1978, D. H. Vilt s.n. (MO). mens of the genus Tort ella from North American This species, as Sarconeurum tortelloides Greene, herbaria, as well as material from the southern was first described from Antarctica (Greene et al. hemisphere, Tortella alpi cola , is reconfirmed taxo­ 1970). Robinson (1972) placed the species into the nomically as a distinct species. Some account of its genus Tortella . Zander and Hoe (1979) included a distribution has begun to emerge, at least in North 1997 NOTES 321

America. It is apparently a species of more widespread ca n reach 6 em , T fra gilis 5 ern in height, and both occurrence than previously thought. are typically two or more cm tall. Since T. alpicola is W ith clarification of its di st inctive charac ters . usuall y only one em in height (rarely 1.5 em ), the Tortella alpicola is readil y differenti ated from the former two spec ies can most often be differentiated two other spec ies in the genu s with which it is most by height alone. It is when specimens of anyone of oft en co nfused: T. tortuosa (Hedw.) Limpr. and T. the thre e species are only one em in height or less fragili s, which are usually robust : T tortuosa that the real confusion arises between them ;

Key to North American specimens of Torre/la tortuosa, T. fragilis and 1: aipicola when stems are only one em high: . I. Leaf apices not deciduous (youngest leaves at the stem apex not propaguloid) with nearly subulate apices composed of more than fivecells; leaf cells II- I I 11m wide Tor/ella tortu osa , I. Leaf apices deciduous (youngest leaves at the stem apex with deciduous, propaguloid modifications) the apices mucronate with fewer cells, occasionally to fivecells, leafcells 9- 13(-14) J.I m wide 2. 2. Propaguloid apices of youngest leaves in one straight segment without articulations, with smooth, straight margins bordered by elongate, smooth cells; stems usually red-tomentose with fine, conspicuous tomentum ...... Tortella fr agilis. 2. Propaguloid apices of youngest leaves articulated into several "barrel-shaped" segments, with crenulated-papillose margins of quadrate cells; stems withouttomentum. or rhizoids sparse and coarse Tortella alpicola.

Th ere are other less definite distinctions. such as genus Tortella . There are also occasional low and the snowy white hyaline basal ce lls in Tortella a/pi­ distant teeth on the upper lea f margin. cola, while these regions in the other two taxa have Specimens of Tortella alpicola see n to date are a yellowish tint. Both Tortella tortuosa and T. f rag­ associated with shaded or exposed. wet or dr y ilis have con spicuous red tom entum, but T. alpicola rocks, crevices and ledges of granite. qu artzite, doe s not. The largest specimens of T. alpicola seen sandstone and calcareous outcrops on cliffs and in tend to be more rhizoidiferous than is typical, wh ile canyons. but also in wet. mesic tundra in the Yukon s ma ll variants of 1'. f ra g ilis te nd to ha ve less at elevations between 900 and 3300 m. In addition tom ent um, but at least so me stem s can be found in to its Canadian range. its general distributio n so far a collection th at are red-to mento se. T he clear, noted is North America in Alaska, Arizona , smoo th elongate cells on the upper margin of espe­ C o lorad o . Idaho, Montana, Nebraska, Uta h, cially young leave s of Tortella fra gilis is a critical Wyoming; South America in Colo mbi a, Asia in character in distinguishin g this species from both T. India, Paci fic Islands in Haw aii. an d Antarctica alpicola and for ms of T. tortuosa with which it may (Alexandra Island ). be co nfused (Zander, manu script in preparation). That a peculiar species of Tortella occurs in our A mo ng th e spec ime ns seen from the so uthern flora has shown itself in the literature in various hemisphere, a tiny example of T. f ragilis fro m ways. Tortella nitida (Lindb.) Broth. was ascribed Ca mpbell Island in Antarc tica proved diffi cu lt to to the North Ame rican flo ra by Haring (19 38) and differentiate fro m T. alp ico /a, mainl y becau se of Flowers ( 1973). Th is ha s been suggested to be a the apparent absence of any propaguliferous leaf varie ty of Tortella tortu osa by Crum and Anderson apex modifications of any distin ction . Alth ough the ( 198 1), who also state that materi al cited as T. niti­ apices of the lea ves were very narro w and decidu­ da by Haring was in fac t either T. f ragilis or T. tor­ ous (fragile), most of them possessed the clear bor­ tuosa. Material from Utah, de scribed by Flowe rs, der of thicker- wall ed elongate cells and, with dili­ does not appea r to have been see n by them. ge nce, one or two apices co uld be found with thick ­ Tortella nitida , a Euro pea n spec ies, ho we ver , ened modificat ion s. Th e lar ger average leaf cells of has basal cell s gradua lly, not abruptly, differentiat­ so me of these redu ced specime ns also duplicate the ed from the laminal cell s, a shining cos ta on the typ ical size of ce lls of T. alpi cola . Th e spec ime n dor sal leaf surface with no di st in ct ive subu late was, however, coarse, with blackened tomentum at propagul oid leaf apex, and leaf cells to 10 urn wide. the very base. The marginal border has proved criti­ Tortella nitid a , by reason of its basal ce lls and cal in naming similar specimens of T. frag ilis. other charac ters, has been thought to possibl y be a Actually, the spe cies that is mo st similar to T. fragile- lea ved form of Trichostomum crispulum (R. alpieola, eve n to the barrel -shaped seg mentatio n in Za nde r, personal co mmunicatio n). North Ame rican th e apic es of the yo ung lea ves, is sma ll, narr ow ­ mater ial exa mined for the present study that was lea ved form s of Triehostomum tenuirostre (Hoo k. nam ed Tortella nitida is var iou sly T. alpie ola , T. & T ayl. ) Lin db . Sp ecimen s of th e latt er species , fragilis, 1'. tortu osa and Trichostomum tenuirostre, howe ver, ar e re adily sepa ra te d by th e thi ck er­ with Tortella alpieola spec ime ns predominating in walled ce lls on the leaf margins at the base , whose the western United St ates of Utah and Colorado . cell s do not form the characteristic vee-s hape of the Tortella alpicola differs from T. nitida immediately 322 THE CA NADIAN FI ELD-NATURALIST Vol. III

in th e brightly di st in ct , clear basal cells set o ff Crum, H. A., and L. E. Anderson. 198 1. of fr om th e g reen lamin al cells as well as th e o ther Eastern North A merica . 2 Vol um es. Co lumbia c haracte rs mentioned. One sp ecimen cited as T. University Press. New York. Dixon, H. N. 1930. Addition s \0 the moss flora of the nit id a from A laska p roved to be Tr ich ost om um North-Western Himalayas. Annales Bryologici I II: tenu irostre, di st inguished by th e basal cells not in a 5 1-70. vee - shape . b u lging-mam mi l lose cells, and tin y Eckel, P. M. 199 1. Torte lla tortelloide s (Mus c i: papill ae often a rranged in two parall e l rows over Pottiaceac) new to North America. Bryologist 94: 84-87. each lumen. Flowers, S. 1973 . Mosses: Utah and the West. Brigham Specimens of T. alpico la fro m Nort h A merica Young University Press, Provo. Utah. have also been calle d T. rigens N . Al berts, (Weber Greene, S. W., D. M. Greene, 1'. D. Brown, and J. M. Pacey. 1970. Antarctic mo ss flora . I . Th e genera 1973) d ue to the wi de leaf cells (ca. 14 pm ), but Andreaea, Poh li a, Poly tric h um , Psil opilu m, an d pl ants of th at species in Sweden arc lon ger, 1.5- 3 Sarconeurum. British Antarctic Survey Science Report em hi gh. a nd lack a p ical p ropagulae (.fee aim 64: 1-11 8. Nyholm 1989). Exam inatio n of m at erial of th is Haring, I. M. 1938. Tortella. In Moss Flora of North s pecies from S wede n sho wel l T. rigens to ha ve a Am erica. Edi l ed by A . 1. Gr out, 1(3): 165- 170 . stro ng morphological sim ilarity to T. [r ag ilis and to Newfane, Venn onl. be different in m an y characters from T. alpicola, Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herhariorum . Pan I . Edition 8. New York es pec ially in leaf shape and co lo r. Botanical Garden, Bronx. New York. HyvoencnvI. 1991. Tar/ella [ragilis () repon ­ Acknowledgments ed for Southern South Am erica . Br yol o gi st 94 : I thank St ewart Scott, Depa rt men t of Geog raphy, 4 16-418. U niversity of Calgary , fo r providing me with a col­ Nyholm, E. 1989. Ill ustrated Flora of Nordi c Mosses. Fasc. 2. POlliaceae-Splachnaceae-Sc histostegaceae. lection of undetermined s peci mens to exa mine, Nordic Bryological Socie ty. Cope nhagen . Denmark. collected by C. D. Bird an d his associates. Robinson, II. E. 1972. Observations on the origin and I th ank th e c u ra to rs o f bryop hytes a t A LTA, of the Antarctic moss flora. Pages 163-177 in C ANM , C HR, COLO, H, MO, and N Y fo r all o w ­ Antarctic Terrestrial Biology. Edited by G. A. Llano . i n g the examin atio n of s pecimen s fro m the i r Antarctic Research Series 20. American Physical Union. herbaria. I am grateful to Richa rd Zander, w ho per­ .Washington, D.C. Weber, W. A. 1973. Guide to the mosses of Colorado. mi tted me access to his un pub lished m a nu script Institute of Arctic and Alpine Research, Univers ity of d iscussing the genus Ta rte // a i n Arc t ic Nort h Colorado, Occasional Paper Numbe r 6. A m e r ic a , and w ho re v ie w ed th e pre sent ·pa pe r. Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of Outs id e reviewe rs c ontributed s u bsta n tially to harsh env ironme nts. Bulletin of the Buffalo Socie ty of im pro ve me nts in the ma nu script. Natural Sciences 32: i-vi + 1-378. Zander, R. H., and W. .I. Hoc. 1979. Geographic dis­ junctio n and heterophylly in To rtel la [ragilis var. Literature Cited tort elloides (=Sa rcan eurum tortelloides) . Bryologist Churchill, S. P., and E. L. Linares C. 1995. Prodromus 82: 84-87. Bryologiae Novo- Granatensis: Introduccion a la Flora de Musgos de Colombia. Partes 1 y 2. Biblioteca "Jose Received 12 August 1996 Jeronimo Triana" 12. Santafe de Bogota. Accepted 29 October 1996