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University of Colorado, BoulderCU ScholarSeries in BiologyEcology & Evolutionary BiologySummer 5-1-1961Studies of Colorado bryophytesWilliam A. WeberFollow this and additional works at: http://scholar.colorado.edu/sbioRecommended CitationWeber, William A., "Studies of Colorado bryophytes" (1961). Series in Biology. 20.http://scholar.colorado.edu/sbio/20This Article is brought to you for free and open access by Ecology & Evolutionary Biology at CU Scholar. It has been accepted for inclusion in Series inBiology by an authorized administrator of CU Scholar. For more information, please contact cuscholaradmin@colorado.edu.

STUDIES OF COLORADO BRYOPHYTES1ByW illia mA.W eber2The bryophyte flora of the Southern Rocky Mountains and ColoradoPlateaus region is both rich in species and interesting from a phytogeographical standpoint. It draws from the arctic or boreal flora, from the dis junct floras of the isolated mountain masses of the northern continents, andfrom an ancient semi-desert flora which is now highly disjunct and limited tothe major desert areas of the world. Thus the great majority of our speciesin Colorado are also found in Eurasia.The bryophyte flora may be said to be distinctly “ continental” in com position because our climate is generally one of extremes in which the mesichabitat is uncommon. Thus, our flora lacks many of the species which are char acteristic, on the one hand, of the eastern deciduous forest and, on the other,of the western Coastal elements of the Cascade region. Nevertheless, a few ofthese species do occur along the east flank of the Front Range adjacent tothe Great Plains because of local topography and weather conditions com bining to produce an unusually cool, moist climatic regime in the foothill can yons. Much of the excitement of bryological work in Colorado comes in thediscovery of these unexpected rare species inhabiting the foothill canyons.The alpine areas are also of great bryological interest. In those choice lo calities above timberline where moisture levels remain continuously highthroughout the year and a well-developed “ moss-tundra” exists, several char acteristic Arctic-alpine species occur at their only stations in the contiguousUnited States. When one does field work in the Colorado alpine he enjoys avicarious experience with the moss flora of Scandinavia, the Swiss Alps and theArctic regions of either hemisphere, for most of the species he encounters arefound in all of these regions.The bryophytes of the drier, semi-desert areas of our southwestern section,while fewer in numbers of species, nevertheless have their special interest.Only a token number are actually restricted to the American Southwest, whilethe great majority occur in other desert regions, particularly the area of theMediterranean Sea, the Near and Middle East.This interesting combination of geographical elements in the Coloradobryophyte flora suggests that the student of these plants must try to acquirea working knowledge of the world’s literature and species. Here is no roomfor narrow provincialism.1 Andreaeaceae, Aulacomniaceae, Cryphaeaceae, Fabroniaceae, Fissidentaceae, Meesiaceae, Neckeraceae,Orthotrichaceae, Polytriehaceae, Splachnaceae, Tetraphidaceae.2 Associate Professor of Biology and Curator o f the Herbarium, University o f Colorado Museum.27

28UNIVERSITY OF COLORADO STUDIESDr. Geneva Sayre, now a professor at Russell Sage College, Troy, NewYork, devoted a great deal of time and effort to a study of the Colorado MossFlora. Her Ph.D. thesis (1938) brought together all that was then known onthe subject. Dr. Sayre also was instrumental in establishing the bryophyteherbarium at the University of Colorado. Her work was a pioneering effort,undertaken at a time when collections of Colorado mosses were very meagre.The thesis preceded by several years the publication of certain portions ofA. J. Grout’s great work on American mosses (1928-1939) so that her treat ments of the Bryaceae and Pottiaceae, for example, could not take advantageof his later findings. Nevertheless, it has been immeasurably useful to have herthesis before me in my studies of the moss flora inasmuch as Dr. Sayre didbring together the known records of the species, both published and un published.Our knowledge of the other great group of bryophytes, the Hepaticae orLiverworts, is extremely fragmentary. Aside from a few lists published inpapers growing out of the Hayden and other historic surveys the only paperdealing with a survey of Colorado hepatics is that of Dr. Alexander W. Evans(1915) which listed only 41 species, most of them from only one or two col lections.I have planned for some years to prepare a revision which would bringup-to-date the work of Dr. Sayre on the mosses and of Dr. Evans on thehepatics. But publication of the entire group at this time would be highlypremature. Several reasons argue for piecemeal treatment. First, those generawhich contain only a few easily recognized species whose distribution is al ready well understood can be disposed of without delay. Many of these speciesare of interest to ecologists, who have an immediate need for information ontheir identification and habits. Second, with more active collection going on,our knowledge of some of the rare alpine and desert species literally grows dayby day; these groups may reach the stage of publication within a very fewyears. But the more difficult or critical groups, such as Bryum, Hypnum,Barbula, and the like, require intensive monographic study, very thoroughcollection, and field observation as well as the co-operation of specialists.These groups may face a long delay in publication.I therefore balance between the desire to fulfill Dr. Sayre’s intention topublish a complete Colorado Moss Flora and the need to make available assoon as possible whatever I can of this flora to those amateur and professionalbotanists who need to know the bryophytes. In choosing the second course westand to gain the interest and enthusiasm of new students of bryology, whowill, in the long run, contribute significantly to the final result.The present series of papers will employ keys, citations of specimens for therecord, and brief discussions pertinent to recognition of the species and their

29STUDIES OF COLORADO BRYOPHYTEShabitats. Excellent technical descriptions and illustrations of all of the NorthAmerican mosses are now generally available in the works of Grout (1903 1939), and the work of Schuster (1953) provides adequate technical aid forthe liverworts. The student should have in his working library the excellentguides of Conard (1956) and Watson (1955). A variety of source-booksis indispensable for the beginner as well as the professional in the field ofbryology.I gratefully acknowledge the generous help and advice of all of my bryological colleagues. Individual credit is given under the citation of specimens.Unless otherwise specified, the collections cited are in the University of Colo rado Herbarium. For convenience of reference, the families are arranged inalphabetical rather than phylogenetic sequence.In the interest of brevity of citation, I am adopting code initials for thecounties of Colorado, after those of the Smithsonian Institution River BasinSurveys. The key to the code initials is as erChaffeeCheyenneClear sDouglasEagleElbertEl sonJeffersonKiowaKit CarsonLakeLa PlataLarimerLas OteroOurayParkPhillipsPitkinProwersPuebloRio BlancoRio GrandeRouttSaguacheSan JuanSan is entire family is characterized by the peculiar dehiscence of the capsule,which splits into four valves with the valves remaining attached at top and

30UNIVERSITY OF COLORADO STUDIESbottom, making a sort of paper-lantern structure unique in the mosses. Ourspecies is strictly alpine, forming tufts, looser than those of most Grimmiaspecies, on siliceous rocks above timberline. The tufts resemble those of Grim mia, but the individual stems are very slender, totally black with sometimesa brownish tinge, and the leaves lack hair-points. The leaves of our only spe cies, A. rupestris, lack a costa.A n d re a e a r u p e s tr isHedw.—Black Moss; Soot Mossb l : on large boulder, west end of the uppermost of the Green Lakes, T. 1 N.,R. 74 W., Boulder Watershed, north of Kiowa Peak, 12,000 ft., Weber 8596.cc: base of large boulder, near the water, inlet of Summit Lake, 12,700 ft.,Mt. Evans, Weber & Jones 8534. e p : Pikes Peak, along the Cog Railway,above 13,000 ft., Holzinger s.n. l r : Rocky Mt. Nat. Park, on vertical wall ofcirque below Lake of Glass, very abundant, 10,000-11,000 ft., Weber &Pontecorvo 9692, Krypt. Exs. Vindob. No. 4153.AULACOMNIACEAEWe have the single genus, Aulacomnium. In Scandinavia these plants arecalled “ Reffelmossor” , the term ‘reffel’ rifling, possibly alluding to thecapsule which is grooved or longitudinally striate. Aulacommium is an acro carpous moss with lanceolate or lance-ovate, decurrent leaves; the midrib isprominent and ends just below the leaf-tip; the margins of the leaves arenarrowly revolute at least below the apex, and the leaf-cells are small(5-15u), isodiametric and papillose, mostly with one papilla over the cellcavity. The leaves may be entire or irregularly denticulate.Aulacomnium. is rarely found fruiting in our area. Instead, the vegetativestems often bear stalks (pseudopodia) which bear clusters of gemmae. Theseta is elongate and the capsule is slightly curved, ascending or almost hori zontal, with a striate or furrowed wall.Keyto t h eS p e c ie sla. Leaves small, mostly less than 1.5 mm. long; basal leaf cells green, not distinctlydifferent from the upper, unistratose; stems characteristically with a terminalpseudopodium bearing a ball-like cluster of few-celled, fusiform gemmae; habitaton rotting w ood . Aulacomnium androgynumlb . Leaves larger, from 2 to 4 mm. long; basal leaf-cells yellow-brown, thick-walled,often enlarged, bistratose; pseudopodia when present naked or bearing relativelyfew green, minute but leaf-like ecostate gemmae in an erect cluster; habitat bogsand slow streams.Aulacomnium palustre

STUDIES OF COLORADO BRYOPHYTES31AULACOMNIUMA u la c o m n iu mandeogynum(Hedw.) Schwaegr.b l : on decaying wood of cut stumps of Pseudotsuga, trail between ScoutCabin and Kossler’s Lake, west base of Green Mountain, S.W. of Boulder,7,000 ft., Weber 4511.Aulacomnium androgynum must be more abundant than our single recordwould suggest. Watson (1955) states that although its typical habitat isrotten wood, it also may occur on humus-rich soil of banks and occasionallyon sandstone rocks. In our area it would be most unusual to expect the speciesto occur on rock because of the arid climatic regime.A u la c o m n iu mp a lu s tr e(Hedw.) Schwaegr.b l : 11 mi. N. of Nederland, 9,500 ft., Pursell 3134', University Camp, 9,900ft., Livingston 108. c f : St. Elmo, 10,000 ft., Kiener 6738. d a : Devil’s HeadMt., 8,000 ft., Weber 7754- g f : Trappers Lake to Coffin Lake, 9,700 ft.,Shushan B-S02. g a : South Fork Columbine Cr., 5 mi. S.E. of Grand Lake,9,500 ft., Douglass B-621. g l : near Tolland, Grout s.n. g n : Elko Basin toMexican Cut, 10,000 ft., Weber 9228; Gothic, 9,800 ft., Sayre 589; Mt. Car bon, Tidestrom 3485. j a : 3 mi. N. of Willow Cr. Pass, 9,000 ft., Shushan 193;Big Creek Lake, 9,200 ft., Shushan 28. l k : Twin Lakes, Conard 451. l r :Moons Ranch, Baker 34 (as A. androgynum); Chambers Lake, Crandall 36.p a : 11.3 mi. E. of Jefferson on road to Lost Creek Park, 10,000 ft., Shushan83. s t : Monte Cristo Creek, N. of Hoosier Pass, 10,800 ft., Weber s.n.;Breckenridge, 10,000 ft., Holzinger 35.This species is one of our most abundant mosses, occurring in almost everybog area in the middle altitudes of the mountains. In its typical form, Aula comnium palustre is very easily recognized by its yellow-green, loosely spread ing upper leaf-growth and very densely shaggy-brown-tomentose lower stems.In the alpine, however, an unusual type occurs which merits further detaileddiscussion below.A u la c o m n iu mp a lu s tr evar.im b r ic a t u mB.S.G.cc: shore of Summit Lake, Mt. Evans, 12,800 ft., Weber B -1401 (Crypt.Exsic. Vindobonensis No. 4160), Weber & Jones 8527, 8541, Weber B-218,9267; 12,500 foot level along trail from Stevens Mine, 11,200 ft., to summit ofGray’s Peak, 14,250 ft., Weber 5642.Some time ago I reported this (Weber 1952) as the Arctic and easternAmerican alpine Aulacomnium turgidum (Wahl.) Schwaegr. Morphologically

32UNIVERSITY OF COLORADO STUDIESour material stands nearer A. turgidum than A. palustre, but the leaf-shapevaries in the direction of palustre to such an extent that one would hesitate toassign the plants unequivocally to turgidum.The general aspect of var. imbricatum is like that of A. turgidum. Thestems are stout and loosely adherent to each other because of the denselycrowded imbricate leaves and a general absence of long-protruding stemtomentum. The stem tomentum is present, but is mostly hidden in the leafaxils. In fact, the tomentum is hardly apparent even on the older growth.The stems form erect sods on wet hummocks which may be saturated withmoisture through most of the growing season. For the most part the leaves arebroad, blunt, rounded at the apex, not at all denticulate, cucullate, and hardlycrispate. However, on some of the stems the terminal growth has more at tenuate leaves which are slightly crisped as in A. palustre.Wallace (1950) commented on the difficu

The bryophyte flora of the Southern Rocky Mountains and Colorado Plateaus region is both rich in species and interesting from a phytogeo- graphical standpoint. It draws from the arctic or boreal flora, from the dis junct floras of the isolated mountain masses of the northern continents, and from an ancient semi-desert flora which is now highly disjunct and limited to the major desert areas .

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