Taconite 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.
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THE CANADIAN FIELD-NATURALIST
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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.
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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 alpic ola 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. IOrtuo sa 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 material of the Geography Department of the University o f Calgary (tho se not s e n t to the Provincial Museum of Alberta in Edmonton in 1995 and examined subsequent to that year), specimens of Tort ella alpicola Dix . c ame to light. The North American distribution of this species, due to taxo nom ic difficulties, has not yet been adequately docu mented. The University of Calgary specimens, and three others seen from Canada at other herbaria, rep resent the pre sent known distribution of the species in that country (herbarium code throughout follow s that des ignated by Holmgren et al. 1990) :
Tortella alpicola : Alberta: Red Deer River Watershed, provincial campsite at j unc tion of James River and Will son Cr eek on Coch rane -Nordegg Forestry Trunk Ro ad. approx. 5 J048' N, 115°03'W, elev. 4400 feet. June 23. 1961, C. D. Bird 6249 ; with En calypta vulgaris (Geography Dep artment , University of Cal gary) : Bow River Watershed, 3 miles WSW of Exshaw, SWI7-24-9-W5, 51°03 ' N, 115°13'W, elev. 4300 ft., Upper Foothills Forest, north-facing slope with steep limestone expo sures, on May 19, 1976, Glennis Lewis no. 36, with Encalypta rhaptocarpa and Gymnoslomum aerugino sum (BUF); Jasper National Park, Dryas drummondii flat 2-3 miles below Athabaska Glacier, July 23, 1959, W. A. Weber B-2278 (COLO). Northwest Territories: Ellesmere Island , lake ca st of Taconite Inlet, 82°52'N, 78°05'W, cliff of metamorphic rock, 50 rn, May 21,1969, G. Brassard 4 136 (CANM). Yukon Territory: Selwyn Mountains. Itsi Range. North end of unnamed lake , south of Fuller Lake , 62-57 'N, 130--09'W. 1600 m, in mesic Betula pumila-Vaccinium uliginosum dominated tundra on cast and north facing slopes, siliceous rock outcrops of granite and slate pre sent, with small amounts of calcareous rock cropping
out, August 10, 1978, D. H. Vilt s.n. (MO). This species, as Sarconeurum tortelloides Greene, was first described from Antarctica (Greene et al. 1970). Robinson (1972) placed the species into the genus Tortella . Zander and Hoe (1979) included a
specimen from Hawaii in the distribution under the name Tortella fragilis val. tortelloides (Greene) Zand. & Hoe, as Zander and Hoe considered it only marginally distinct from Tort ella fragilis (Hook. & Wils in Drumm.) Limpr. After study of material of the genus Tortella from throughout Arctic North America, Zander (m anuscript in preparation) con cluded that the variety was a good species, as origi nally determined by Robinson . Following Zander's decision, Eckel (1991) pub lished the discovery of the species in the contermi nous United States from the sta te of Wyoming , including reference to a station in the Northwest Territories of Canada. The taxon was discussed again by Hyvoenen (1991), who indicated, based on his examination of specimens of Tortella fragilis from the southern hemisphere, that Tortella tortel loides should be plac ed under Tortella fragilis as a variety again . My examination of specimens at H from Ti erra del Fuego collected by H . Roivainen, some of which were annotated by Hyvoenen , has shown all of the se specimens to be Tortella fragilis val. [ragili s. None of them were what is indicated here as Tortella alpicola. Other specimens at CHR, H , NY and ALTA of both Tortella [ragili s and T. tortuosa collected in the southern hemi sphere were all either correctly determined or were species other than T. alp icola. So far, the only known southern hemisphere s ta tio ns for the latter species are in Antarctica. The species has been found in Colombia (Churchill and Linares 1995), and may prove to be a member of the Andean flora . Zander (1993) reported an earlier name for Tortella tortelloides based on the type of T. alpicola Dixon (1930), collected from a station in the Himalayas.
As a result of my examination of numerous speci mens of the genus Tort ella from North American herbaria, a s well as material from the southern hemisphere, Tortella alpi cola , is reconfirmed taxo nomically as a distinct spe cies. Some account of its distribution has begun to emerge, at lea st in North
1997
NOTES
Am erica. It is apparently a species of more widespread occurrence than previously thought. W ith clarification o f its di st inctive c harac te rs . Tortella alpi cola is readil y d iffer enti ated from the two other spec ies in the genu s with whi ch it is most oft en co nfused: T. tortuosa (Hedw.) Limpr. and T. fragili s , which are usually robu st plants: T to rtuosa
321
ca n reach 6 em , T fra gilis 5 ern in height, and both are typi call y 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 speci mens 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 five cells; 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
2. mucronate with fewer cells, occasionally to five cells, leaf cells 9- 13(- 14) J.Im wide 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 Tortella alpicola. margins of quadrate cells; stems withouttomentum. or rhizoids sparse and coarse Th ere are other less definite distinct ions. such as the sno wy white hyaline basal ce lls in Tortella a/pi cola , wh ile these regions in the other two tax a have a ye llowish tint. Both Tortella tortuosa and T. f rag ilis have con spicuous red tom entum, but T. alpicola doe s not. Th e large st specime ns of T. alpicola seen tend to be more rhizoidiferou s than is typical, wh ile s m a 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 c oll e ction th at are re d - to men to se . T he clear, smoo th elon gate cells on the upper marg in of espe cially yo ung leave s of Tort ella fra gilis is a critical ch aracter in distin gu ishin g thi s spec ies from both T. alpicola and for ms of T. to rtuosa with which it may be co nfused (Za nde r, manu script in prep aration ). A mo ng th e spec ime ns see n from the so uthern hem isphere, a tiny e xampl e o f T. f ra g ilis fro m Ca mpbe ll Island in A ntarc tica pro ved diffi cu lt to differ enti ate fro m T. alp ico /a, mainl y becau se o f the appa rent abse nce of any propaguli ferou s leaf apex mod ification s of any distin cti on . Alth ou gh the ap ices of the lea ves we re very narro w and dec idu ou s (fragile), most of them posse ssed the clear bor der of th icker- wall ed elongate ce lls and, with dili ge 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 o f these redu ced specime ns also duplicate the typ ic al size of ce lls of T. alpi cola . Th e spec ime n wa s, however, coar se, with blackened tomentum at the very base. The marginal border has proved criti ca l in naming similar speci me ns of T. frag ilis. Actually, th e spe ci e s th at is mo st si m ilar to T. alp ieola, eve n to the barr el -shaped seg me ntatio n in th e apic e s of th e yo ung lea ves, is sma ll , narr ow lea ved form s of Trieh ostomum tenuirostre (Hoo k. & T ayl. ) Lin db . Sp ecimen s of th e latt er species , how e ver, ar e re adil y se pa 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 characteristic 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 lora d o . Idaho, Montana, Nebras ka, U ta h , W yoming ; So uth America in C o lo mbi a, A sia in India, Pa ci fi c Is land s in Haw a ii. an d Antarc tic a (Alexa ndra Island ). That a peculiar species of Tortella occurs in our flora has sho wn itself in the literature in va rio us ways. Tortella nitida (Lindb.) Broth . was as cribed to the North Ame rican flo ra by Haring (19 38) and Flowe rs ( 1973) . Th is ha s bee n suggested to be a varie ty of Tort ella tortu osa by Crum and Anderson ( 198 1), wh o also state th at materi al cited as T. niti da by Haring was in fac t e ither T. f rag ilis or T. tor tuosa . Material from Utah, de scr ibed by F lowe rs, doe s not appea r to have been see n by them. Tortella nit id a , a E uro pea n spec ie s , ho we ver , has basal cell s gradua lly, not abruptly, d ifferentiat e d from the lamin al cell s, a shining cos ta on the d or sal le af surface with no di st in ct i ve s ubu late propagul oid leaf ape x, and leaf cells to 10 urn wid e. Tortella nit id a , by re ason o f it s b asal ce lls and other charac ters, has been thoug ht to possibl y be a fra gile- lea ved fo rm of Trichostomum crispulum (R. Za nde r, per sonal co mmunic atio n). North Ame rican mater ial exa mine d for th e pr e sent study th at was nam ed To rtella nitida is var iou sly T. alpie ola , T. fragilis, 1'. tortu osa and Trichostomum tenuirostre, with Tortella alp ieola spec ime ns predominating in the we stern Uni te d St ate s o f Uta h and Co lo rado . Tortella alpicola differs fro m T. nitida immediately
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THE CA NADIAN FI ELD-NATURALIST
in th e bri ghtl y di st in ct , clear bas al ce lls se t o ff fr om th e g reen lam in al ce lls as w ell as th e o ther c harac te rs m ent ion ed . One sp eci me n cited as T. nit id a f rom A las ka p ro v ed to be Tr ich ost om um tenu irostre , di st inguished by th e basal cells no t in a vee - sh ape . b u lgi ng- ma m mi l lose ce lls , an d t in y p a p ill ae often a rranged in two p arall e l rows over each lumen. S p e c i me ns of T. alpico la fro m No rt h A me rica have also been ca lle d T. rigens N . Al berts, ( We be r 1973 ) d ue to the wi de leaf ce lls (ca. 14 pm ), but pl ants of th at spec ie s in Sweden a rc lon ger, 1.5- 3 e m hi gh. a n d la ck a p ica l p rop a gul a e (.fee aim Nyh olm 1989). Exa m i natio n of m at eri al of th is s pecies from S w e de n s ho w el l T. rigen s 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 sha pe and co lo r. Acknowledgments I thank St ewart S c ott, D e pa rt m en t of G eo g raph y , U niversi ty of Ca lga ry , fo r providing m e w it h a co l le cti on o f undetermin ed s p e ci 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 bryo p hytes a t A LTA, C A N M , C H R , COLO, H, MO , a nd 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 he rbaria. I am gratefu l to R ic ha rd Zander, w ho pe r mi tted m e acces s to his un pu b lishe d m a nu s cript d i sc uss i ng th e ge nus Ta rte // a i n Arc t ic Nor t h A m e r ic a , an d w ho r e v ie w ed th e pre s en t ·p a p e r. Ou ts id e rev iew e 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 Ch urchill, S. P., and E. L. Linares C. 1995. Prodromus Bryologiae Novo- Granatensis: Introd uccion a la Flora de Musgos de Colombia. Partes 1 y 2. Bibl ioteca "Jose Jeronimo Tri ana" 12. Santafe de Bogota.
Vol. III
Crum, H. A., a nd L. E. Anderson. 198 1. Mosses of Eastern Nort h A meri ca . 2 Vol um e s. Co lumbia University Press. New York. Dixon, H. N. 1930. Addi tion s \0 the moss flora of the No rth-Western Himala yas. Annales Br yo logic i I II : 5 1-70. Eckel, P. M. 199 1. Tort e lla torte lloi de s (M us c i: Potti aceac) 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 flo ra . I . Th e gene ra Andreaea, Poh li a, Pol y t r ic h um , Psil op ilu m , an d Sarconeurum. British Antarcti c Surv ey Science Report 64: 1-11 8. Haring, I. M. 1938. To rtella. In Moss Fl or a of No rth Am eri c a . 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 , Bron x. New York . HyvoencnvI. 199 1. Tar/ella [ragilis (Pottiaceae) repon e d for S outh ern S outh Am eri c a . 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 ac hnaceae-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. America n 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, Occa sional Paper Numbe r 6. Zander, R. H. 1993. Genera of the Pottiaceae: Mos ses of hars h env ironme nts. Bull etin of the Buffalo Socie ty of Natural Sciences 32: i-vi + 1-378. Zander, R. H., and W. .I. Hoc. 1979 . Geographic d is junctio n a nd heterop hyll y in To rtel la [ra gilis var. tort el loides (=Sa rcan eurum tortelloides) . Bryol ogist 82: 84-87. Received 12 August 1996 Accepted 29 Octo ber 1996
