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320 THE CANADIAN FIELD-NATURALIST Vol. 111

The Moss Tortella 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.

1 Eckel, r. 1997. The moss Tortella alpicola Dix, new to Alberta and the Yukon Territory with a discussion of its range and . 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 Tortella fragilis val. tortelloides (Greene) University o f Calgary (those not se n t 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 origi­mented. 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) pub­resent the present known distribution of the species lished the discovery of the species in the contermi­in 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 discussedTortella alpicola :

again by Hyvoenen (1991), who indicated, based on Alberta: Red Deer River Watershed, provincial campsite at his examination of specimens of Tortella fragilisj unc tion of James River and Will son Creek on from the southern hemisphere, that Tortella tortel­Coch rane -Nordegg Forestry Trunk Road. approx.

5 J048' 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.

all either correctly determined or were species other Weber B-2278 (COLO). than T. alp icola. So far, the only known southern Northwest Territories: Ellesmere Island , lake cast of hemisphere s ta tions for the latter species are inTaconite Inlet, 82°52'N, 78°05'W, cliff of metamorphic

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 taxo­1970). 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 occurrence than previously thought.

W ith clarification of its di st inctive charac te rs . Tortella alpicola is readil y d ifferenti ated from the two other spec ies in the genu s with which it is most oft en co nfused: T. tortuosa (Hedw.) Limpr. and T. fragili s , which are usually robust plants: T to rtuosa

ca n reach 6 em , T fra gilis 5 ern in height, and both are typically two or more cm tall. Since T. alpicola is usuall y only one em in he ight (rarely 1.5 em ), the former two spec ies can most often be differentiated by height alone . It is when specimens of anyone of the thre e species are only one em in height or less 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/e lla to rtu 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.Im 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 distinct ions. such as the sno wy white hyaline basal ce lls in Tortella a/pi­cola , wh ile these regions in the other two tax a have a ye llowish tint. Both Tortella tortuosa and T. f rag­ilis have con spicuous red tom entum, but T. alpicola doe s not. The largest specimens of T. alpicola seen tend to be more rhizoidiferous than is typical, wh ile s ma ll vari ants of 1'. f ra g ilis te nd to ha ve less tom ent um, but at least so me stem s can be found in a coll ection th at are red- to men to se . T he clear, smoo th elongate cells on the upper margin of espe ­cially young leave s of Tortella fra gilis is a critical character in distingu ishin g thi s species from both T. alpicola and for ms of T. tortuosa with which it may be co nfused (Za nder, manu script in prep aration).

A mo ng th e spec ime ns seen from the so uthern hem isphere, a tiny exampl e of T. f rag ilis fro m Ca mpbe ll Island in Antarc tica proved diffi cu lt to differenti ate fro m T. alp ico /a, mainl y becau se of the appa rent absence of any propaguli ferou s leaf apex mod ifications of any distin cti on . Alth ough the ap ices of the lea ves were very narro w and decidu­ous (fragile), most of them possessed the clear bor ­der of th icker- wall ed elongate ce lls and, with dili­ge nce, one or two apices co uld be found with thi ck ­ened mod ificat ion s. Th e lar ger average leaf ce lls of so me of these redu ced specime ns also duplicate the typ ical size of ce lls of T. alpi cola . Th e spec ime n was, however, coarse, with blackened tomentum at the very base. The marginal border has proved criti ­ca l in naming similar specimens of T. frag ilis.

Actually, the spe cies that is mo st si milar to T. alpieola, eve n to the barrel -shaped seg mentatio n in th e apic es of the yo ung lea ves, is sma ll , narr ow ­lea ved form s of Triehostomum tenuirostre (Hoo k. & T ayl. ) Lin db . Sp ecimen s of th e latt er species , howe ver, ar e re adil y sepa ra te d by th e thi ck er ­walled ce lls on the leaf margins at the base , wh ose cell s do not form the characteris tic vee-s hape of the

genus Tortella . There are also occasional low and distant teeth on the upper lea f margin.

Specimens of Tortella alpicola see n to date are associated with shaded or exposed. wet or dr y rocks, crevices and ledges of granite. qu artzite, sandstone and calcareous outcrops on cliffs and in canyons. but also in wet. mesic tundra in the Yukon at elevations between 900 and 3300 m. In addition to its Canadian range. its general distributio n so far noted is North America in Alaska, Arizo na , C o lorad o . Idaho, Montana, Nebras ka, Uta h , W yoming ; South America in Colo mbi a, Asia in India, Paci fic Is lands in Haw aii. an d Antarctica (Alexa ndra Island ).

That a peculiar species of Tortella occurs in our flora has shown itself in the literature in va rious ways. Tortella nitida (Lindb.) Broth . was ascribed to the North Ame rican flo ra by Haring (19 38) and Flowers ( 1973) . Th is ha s been suggested to be a varie ty of Tort ella tortu osa by Crum and Anderson ( 198 1), who also state that materi al cited as T. niti­da by Haring was in fac t e ither T. f ragilis or T. tor­tuosa . Material from Utah, de scr ibed by Flowe rs, does not appea r to have been see n by them.

Tortella nit ida , a Euro pea n spec ies , ho we ver , has basal cell s gradua lly, not abruptly, d ifferentiat­ed from the laminal cell s, a shining cos ta on the dor sal leaf surface with no di st in ct ive subu late propagul oid leaf apex, and leaf cells to 10 urn wide. Tortella nit id a , by reason of it s basal ce lls and other charac ters, has been thoug ht to possibl y be a fragile- lea ved form of Trichostomum crispulum (R. Za nde r, per sonal co mmunic atio n). North Ame rican mater ial exa mined for the present study that was nam ed Tortella nitida is var iou sly T. alpie ola , T. fragil is, 1'. tortu osa and Trichostomum tenuirostre, with Tortella alpieola spec ime ns predominating in the western Uni ted St ates of Utah and Colo rado . 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 se t o ff fr om th e g reen lamin al ce lls as well as th e o ther c harac te rs mentioned . One sp eci men cited as T. nit id a f rom A las ka p roved to be Tr ich ost om um tenu irostre , di st inguished by th e basal cells not in a vee - shape . b u lgi ng- mam mi l lose ce lls , and t in y pap ill ae often a rranged in two parall e l rows over each lumen.

S pec imens of T. alpico la fro m Nort h A merica have also been calle d T. rigens N . Al berts, (Weber 1973) d ue to the wi de leaf ce lls (ca. 14 pm ), but pl ants of th at spec ie s in Sweden arc lon ger, 1.5- 3 em hi gh. a nd lack a p ical p rop agul ae (.fee aim Nyholm 1989). Exa m inatio n of m at eri al of th is s pecies from S we de n s ho wel l T. rigens to ha ve a s tro ng morphological sim ilar ity to T. [r ag ilis and to be different in m an y characters from T. alpicola, es pec ia lly in leaf shape and co lo r.

Acknowledgments I thank St ewart Scott , D epa rt m en t of G eog raphy ,

U niversi ty of Calgary , fo r providing m e with a col­lection o f undetermined s peci me ns to ex a mine, collected by C. D. B ird an d h is associates.

I th ank th e c u ra to rs o f bryop hytes a t A LTA, C A N M , C H R, COLO, H, MO, and N Y fo r all o w ­

i n g the ex ami n ati o n of s pec imen s fro m th e i r herbaria. I am gratefu l to R icha rd Zander, w ho per ­mi tted m e access to his un pu b lished m a nu script d i sc uss ing th e ge nus Ta rte // a i n Arc t ic Nor t h A m e r ic a , and w ho re v ie w ed th e pre sen t ·pa pe r. Ou ts id e rev iewe r s c ontributed s u bs ta n tia lly to im pro ve m e nts in the m a nu script.

Literature Cited Churchill, S. P., and E. L. Linares C. 1995. Prodromus

Bryologiae Novo- Granatensis: Introduccion a la Flora de Musgos de Colombia. Partes 1 y 2. Bibl ioteca "Jose Jeronimo Triana" 12. Santafe de Bogota.

Crum, H. A., and L. E. Anderson. 198 1. Mosses of Eastern North A meri ca . 2 Vol um es. Co lumbia University Press . New York.

Dixon, H. N. 1930. Addi tion s \0 the moss flora of the No rth-Western Himalayas. Annales Bryologic i I II : 5 1-70.

Eckel, P. M. 199 1. Torte lla torte lloi de s (M us c i: Pottiaceac) new to North America. Bryologist 94: 84-87.

Flowers, S. 1973 . Mosses: Utah and the West. Brigham Young University Press, Provo. Utah.

Greene, S. W., D. M. Greene, 1'. D. Brown, and J. M. Pacey. 1970 . Antarc tic mo ss flora . I . Th e genera Andreaea, Poh li a, Poly t r ic h um , Psil opilu m, an d Sarconeurum. British Antarctic Survey Science Report 64: 1-11 8.

Haring, I. M. 1938. Tortella . In Moss Flora of No rth Am eri ca . Edi l ed by A . 1. Gr out , 1(3) : 165- 170 . Newfane, Venn onl.

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herhariorum . Pan I . Edition 8. New York Botanical Garden, Bronx. New York .

HyvoencnvI. 1991. Tar/ella [ragilis (Pottiaceae) repon ­e d for Southern South Am eri ca . Br yol o gi st 94 : 4 16-4 18.

Nyholm, E. 1989. Ill ustrated Flora of Nordi c Mosses. Fasc . 2. POlli aceae-Spl achnaceae-Sc hi sto stegaceae. Nordic Bryological Socie ty. Cope nhagen . Denmark.

Robinson, II. E. 1972. Observations on the origin and taxonomy of the Antarctic moss flora. Pages 163-177 in Antarctic Terrestrial Biology. Edited by G. A. Llano . Antarctic Research Series 20. American Physical Union. .Washington, D.C.

Weber, W. A. 1973. Guide to the mosses of Colorado. Institute of Arc tic and Alpine Research, Univers ity of Colorado, Occasional Paper Numbe r 6.

Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of harsh env ironme nts. Bulletin of the Buffalo Socie ty of Natural Sciences 32: i-vi + 1-378.

Zander, R. H., and W. .I. Hoc. 1979 . Geographic dis­junctio n and heterophylly in To rtel la [ra gilis var. tort el loides (=Sa rcan eurum tortelloides) . Bryologist 82: 84-87.

Received 12 August 1996 Accepted 29 October 1996