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At the University of Vermont (UVM), we offer a year-long, interdisciplinary, residentially based course for first-year students interested in the natural sciences. This course, entitled “The Geology and Ecology of Lake Champlain,” takes advantage of the University's location between Lake Champlain and the Green Mountains. Although the University offers students introductory-level courses in both geology and ecology, these are high-enrollment lecture-format classes with some field trips. “The Geology and Ecology of Lake Champlain” offers a small group of students a field-based course that seeks to integrate these related disciplines. In the course, the students become active hands-on participants in small research projects. Although UVM's geographic location has certain unique features, we believe that the fundamental approach of this course can be adopted by universities in other locales.

Eight sites of known age were sampled in Glacier Bay National Monument, Alaska, to examine the changes in plant diversity during primary succession in that region. Four strata–trees, tall shrubs, low shrubs—herbs, and bryoid—thalloids–were sampled independently. Data suggested a sequence of wave—like invasions on sites by strata, largely in order of increasing size. Bryoid—thalloids were exceptional in demonstrating a late peak in cover values. In general, diversity of a particular stratum declined during the period in which the stratum dominated the community in terms of foliar cover. Richness (species number) of communities increased rapidly in the first 100 years, then more gradually to reach a maximum in the muskeg steady state. Equitability (evenness of distribution of foliar cover among species) was erratic, but tended to increase with age. After initial rises, three diversity indices showed nearly flat curves with two exceptions: marked decreases in the 30— to 50—year period; and a rise to maximum levels in the final steady state. Total information per unit area calculated from diversity and foliar cover increased in a sigmoid manner with peak information in the final muskeg stage.

The previous catalog of mosses known from Alaska was published in 1902 and included 280 species and varieties. The present enumeration, compiled from published records, lists 601 taxa in 150 genera and 43 families. Bryological research in Alaska has had two periods of development. The first began with small collections made in the mid-nineteenth century, often by members of a general scientific expedition. By 1902 Cardot and Th~riot (1902) were able to list, in their enumeration of the mosses collected by the Harriman Expedition to coastal regions of Alaska, 280 species of mosses based upon their own collections and those of some ten previous reports. The 1913 Kelp Expedition, also primarily a coastal study, added an additional 55 taxa (Holzinger & Frye, 1921). R. S. Williams (1901, 1903, 1912, 1919), in studies primarily centered in the Yukon Territory, added several species. In 1914 Evans was able to report 105 hepatics from Alaska. (Worley (1970) gives a list of liverworts now known from Alaska.) During these early years many collections were made by nonbryologists and were usually from sites seldom far from the coast. A few, such as those by Hazen and Turner (1886) were made in the interior, but mountains and the Arctic were rarely investigated bryologically. From 1921 until the early 1940's there were few publications on Alaskan mosses. The field work of E. Hult~n in 1932 begins the second era of Alaskan bryology. Bartram (1938) reported the mosses collected from the south coastal regions by Hult6n and by W. J. Eyerdam, a long-time amateur collector of plants. It was, however, in the continuing accumulation of vascular-plant specimens from throughout Alaska by Hult~n that Herman Persson found the material for many years of bryophyte investigations. From herbarium sheets Persson located, separated, determined, and reported in a series of papers (many in THE BRYOLOGIST) the bryophytes that had accidentally been pressed along with the intended plants. From the 1940's to the present he also reported (often in joint authorship) the collections of a number of nonbryologists and bryologists from numerous, widespread Alaskan localities. Notable contributors have been E. Hult~n and Frbre E. Lepage who collected the first substantial arctic material, L. D. Stair and his daughter who collected in Yakutat Bay, and several important collections from the interior, especially the mountain ranges, by W. Weber, L. A. Viereck, H. T. Shacklette, E. M. Sherrard, O. Gjaerevoll, and others. Many of these workers have also published their findings separately. Persson also re-examined 1 We thank Dr. W. B. Schofield for advice throughout the preparation of this list and for financial assistance through the National Research Council of Canada. 2 Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada. * Hattori Botanical Laboratory, Obi, Nichinan, Miyazaki Prefecture, Japan. This content downloaded from 207.46.13.111 on Tue, 09 Aug 2016 05:19:39 UTC All use subject to http://about.jstor.org/terms 60 THE BRYOLOGIST [Volume 73 the Alaskan material in the Kindberg herbarium collected by J. Macoun. Other bryological investigations were made by R. Thomas at Iliamna Lake (1952), by A. M. Harvill from various Alaskan sites (1947a, 1947b, 1948, 1950), by Frye and Clark (1946) reporting on material from Attu Island, and by several other workers reporting in a variety of papers, notes, and monographs. During this period the gap in arctic studies was being filled by the research of W. C. Steere. His many trips to the Arctic brought the knowledge of Alaskan arctic bryophytes up to the level of most of the other regions in the state. Although never presenting a complete list of arctic Alaska mosses, Steere reported many new records in his numerous articles on the North American Arctic. Bryological studies in Alaska have now established the major floristic components, but many distributions are incomplete and a number of critical areas, especially certain mountain ranges and some coastal regions, need intensive study. Many species new to the state, and perhaps even to science, are certainly yet to be discovered (for example, the recent discoveries of Crumia deciduidentata (Sharp & Iwatsuki, 1969) and Pterigoneurum arcticum (Steere, 1959). Major collections in the last five years have been made by A. J. Sharp and Z. Iwatsuki in the Aleutian Islands and the Alexander Archipelago and by I. A. Worley in southeastern Alaska. There are also unpublished collections by F. J. Hermann and B. Neiland. Duplicates of some of the Hermann specimens have been distributed. The last published report attempting to list all known Alaskan mosses was made in 1902 (Cardot & Th6riot, 1902). Persson (1949) totaled, but did not list, the species of mosses from Alaska and the Yukon. His count of published and unpublished records of mosses was 27 Sphagna, 265 acrocarpous mosses, and 140 pleurocarpous mosses, making a total of 432 taxa. Since that time there have been numerous additions to the Alaskan flora. With the completion of E. Hult6n's Flora of Alaska and Neighboring Territories (1968), it was felt that an enumeration of the mosses now reported was due. The present list was compiled from the literature and includes 33 Sphagna, 354 acrocarpous mosses, and 214 pleurocarpous mosses-a total of 601 taxa. With few exceptions the synonymy of Crum, Steere and Anderson (1965) has been followed. For the treatment of the Mniaceae the revisions of Koponen (1968) have been utilized. No attempt has been made to verify determinations or species reports. Occasional entries are almost certainly misdeterminations; they are single reports of common species otherwise not known from the northwest Pacific region and have not been recollected in recent times. Examples of these include Polytrichum ohioensis, Ulota hutchinsiae and var. rufescens, and Cirrophyllum illecebrum. However, until the material is examined they have been retained in the list. When a taxon is known from Alaska from only one or two reports a reference is cited following the name of the taxon in the list. Subspecific taxa are a problem because many of the older ones, which have not been reported since the original typification, have never been reduced to synonymy, although it is clear that they should be. In general, we have omitted those which are excluded by Grout (1928--40) and by Crum, Steere, and Anderson (1965). Some names included are dubious as good taxa but they are retained pending further study. In the genus Sphagnum, with one exception, the list does not include varieties or subspecies. Grout, in the Moss Flora of North America (1928-40), gives without specimen citation the only Alaskan report for the following taxa: Polytrichum commune var. perigoniale, P. ohioense, Ditrichum pusillum, Ceratodon purpureus var. conicus, This content downloaded from 207.46.13.111 on Tue, 09 Aug 2016 05:19:39 UTC All use subject to http://about.jstor.org/terms 1970] WORLEY & IWATSUKI: MOSSES OF ALASKA 61 Trematodon ambiguus, Oncophorus wahlenbergii var. compactus, Dicranum muhlenbeckii var. cirratum, Plagiobryum zierii, Amblyodon dealbatus, Anacolia menziesii, Philonotis fontana var. borealis, Zygodon rupestris, Hygrohypnum bestii, and H. luridum var. julaceum. Both Funaria hygrometrica var. arctica and F. microstoma var. obtusifolia appear in the list although Persson (1962a) considers that perhaps these records represent but one arctic taxon. Brotherella roellii has been reported from Kuiu Island by Harvill (1950). It is possible that this is actually an undescribed species of Brotherella known from British Columbia (Schofield, 1968). The Rhacomitrium heterostichum complex in western North America is currently under study by K. Banu of the University of British Columbia; her findings, soon to be reported, have not been incorporated in this paper. The list is presented by families in the order used by Crum, Steere, and Anderson (1965). THE LIST OF SPECIES

Fifteen species and one variety of mosses are added to the checklist of Alaskan mosses. To the recent checklist of the mosses of Alaska by Worley and Iwatsuki (1970) can be added the following taxa which were inadvertently omitted. The majority were reported by Steere (1954, 1965) and were called to the authors' attention by Dr. G. R. Brassard. Each of the entries has been reported but a single time from the state with the exceptions of Andreaea rupestris var. alpestris, Brachythecium asperrimum, and Thamnobryum leibergii which are known from several records. Andreaea rupestris var. alpestris (Thed.) Sharp Anoectangium sendtnerianum B.S.G. (Steere, 1965) Brachythecium asperrimum (Mitt.) Sull. Desmatodon obtusifolius (Schwaegr.) Schimp. (Steere, 1965) Distichium hagenii Ryan ex Philib. (Steere, 1954) Ditrichum zonatum (Brid.) Kindb. (Schofield, 1968) Fissidens arcticus Bryhn (Steere, 1954) Grimmia plagiopodia Hedw. (Steere, 1965) Mielichhoferia mielichhoferi (Funck ex Hook.) Loeske (Schofield, 1959) Orthothecium strictum Lor. (Persson, 1946) Orthotrichum anomalum Hedw.(Williams, 1901) Seligeria campylopoda Kindb. (Steere, 1965) Seligeria diversifolia Lindb. (Steere, 1965) Seligeria pusilla ( Hedw.) B.S.G. (Steere, 1965) Tetradontium brownianum (Dicks.) Schwaegr. (Steere, 1965) Thamnobryum leibergii (Britt.) Nieuwl. These citations increase the total of taxa reported from the state to 617, in 151 genera. We thank Dr. G. R. Brassard for his critical scrutiny of our previous list. Work on the original checklist was supported by financial assistance from the National Research Council of Canada through Dr. W. B. Schofield.

The last published list of all hepatics known from Alaska included 113 species and varieties. The present enumeration, compiled from published records, lists 208 taxa in 66 genera and 35 families. Hepaticology in Alaska has had two periods of development, similar to the history of the study of mosses (Worley & Iwatsuki, 1970). Contributors who reported the early collections, which were primarily made in south coastal regions, include Stephani (1844-47, 1887) and Kurtz (1895), who discussed the material collected by the Krause brothers in the Chilkoot region of southeastern Alaska; Underwood (1882, 1884, 1891), who determined material from several Alaskan collections; Hazen and Turner (1886), who collected in the interior of the Territory; Pearson (1890), who noted several Alaskan records in his list of Canadian Hepaticae; Howe (1897, 1899, 1901), who contributed several new Alaskan records; Coville (1896), who reported material from Yakutat Bay; and Cooley (1892), who collected in the Alexander Archipelago. By the turn of the century A. W. Evans had begun to work on Alaskan hepatics, reporting the collections of the Harriman Alaska Expedition (Evans, 1900), those of Setchell from western Alaska (Evans, 1901), and in a major paper those of the Kelp Expedition (Evans, 1914). In this last report he listed 105 species. During the next 25 years there was a lull in the study of Alaskan liverworts. Evans added some new records, monographic studies occasionally added others, but no large collections were made. In 1934 Brinkman listed only 109 species from Alaska, and was able to add only four additional species by 1938. Several liverworts were reported for the first time from Alaska in the compilation of North American hepatics by Frye and Clark (1937-47). While preparing this flora they were able to study specimens collected by many of the early expeditions and by more recent investigators. These were reported (Clark & Frye, 1942) along with the plea for future collectors to move inland as nearly all Alaskan material was from near the sea. The second phase in the study of Alaskan hepatics has been almost identical to that of the mosses. Most workers and many of the papers from the early 1940's until the present deal with all bryophytes. The most important contributors have been indicated by Worley and Iwatsuki (1970). Although H. Persson never collected in Alaska, his research on the collections of others have been central to the understanding of I thank Drs. Z. Iwatsuki, M. Mizutani, and T. Amakawa for comments, and Dr. W. B. Schofield for guidance throughout the preparation of the list and for financial assistance through the National Research Council of Canada. 2 Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada. This content downloaded from 207.46.13.101 on Thu, 21 Apr 2016 05:46:58 UTC All use subject to http://about.jstor.org/terms 1970] WORLEY: HEPATICAE OF ALASKA 33 Alaskan bryophytes for the past three decades. Three other workers must be especially singled out: Persson (1968) cites E. Hultin, W. C. Steere, and H. T. Schacklette as "the most successful collectors of bryophytes in Alaska and Yukon Territory." The many publications by Persson and others during this period brought the bryological knowledge of the interior and northern parts of Alaska to a similar stage as that of the coastal regions and nearly doubled the number of taxa of liverworts known from the state. Even though he has never worked specifically with Alaskan hepatics, R. Schuster has offered many important taxonomic judgments of taxa present in the Pacific Northwest. Almost all species known from Alaska are (or will be) discussed, either as main entries or as notes, in his flora The Hepaticae and Anthocerotae of North America East of the Hundreth Meridian, the first two volumes having already appeared (Schuster, 1966, 1969). The major components of the Alaskan hepatic flora have now been established, but many problems still exist, some of them requiring intensive and critical study. The number of hepaticologists who have collected in Alaska is small, and the collections have not been extensive. A frequent consequence has been that the mosses have received more attention, both in the field and in herbaria. That there are many species of liverworts yet to be found, especially in the alpine regions, is underlined by the recent discoveries of Takakia ceratophylla (Sharp & Hattori, 1967), Acrobolbus ciliatus (Sharp & Hattori, 1968), Gymnocolea acutiloba, Hybrobiella laxifolia, and Lophozia opacifolia (Persson, 1968), and Haplomitrium hookeri (Worley, 1969). An addition of another 100 species to the present list is certainly to be expected. Major collections in the last five years have been made by A. J. Sharp and Z. Iwatsuki in the Aleutian Islands and the Alexander Archipelago and by I. A. Worley in southeastern Alaska. The most recent enumeration of liverworts known from Alaska was made by Persson in 1949, but it contained no list of species. Persson noted a total of 160 published and unpublished records of hepatics. The present list was compiled from the literature and contains 208 species and varieties in 66 genera and 35 families. Family and generic systematics and, with few exceptions, the species treatment follow Schuster (1966, 1969). For most taxa not yet treated by Schuster the work of Arnell (1956) is followed. The conservative treatment of the Lophoziaceae by Schuster (1969) has been used and the subgenera of the genus Lophozia are indicated within the list. No attempt has been made to verify determinations or species reports. Hence, occasional entries may represent misdeterminations. Persson (1968) emphasizes the need for critical study of several genera in Alaska, namely Calypogeia, Cephaloziella, Lophozia, Plagiochila, and Scapania. To these Marsupella and Frullania can be added. In the list that follows the names of taxa known from Alaska from only one or two reports are indicated by a reference. A few critical discussions in the literature are similarly indicated. The status of the species of Frullania from Alaska is somewhat unclear. Stephani's species F. chilcootiensis from Chilkoot, Alaska, is known only from the type and is poorly described. It is probably only a form of a common coastal species (Frye & Clark, 1937-47). Frullania californica may not be in Alaska (Persson, 1952); F. frasciscana is known from several collections, but has not been reported since the list of Frye and Clark (1937-47); and F. tamarisci is rarely reported, the most recent record also being that of Frye and Clark. Frullania nisquallensis is common and abundant within its range. This content downloaded from 207.46.13.101 on Thu, 21 Apr 2016 05:46:58 UTC All use subject to http://about.jstor.org/terms 34 THE BRYOLOGIST [Volume 73 Calypogeia neesiana has also not been reported recently from Alaska. It is possible that the early reports represent other species. Moerckia hibernica has been considered by some as a synonym of M. flotowiana. This makes certain citations unclear. Persson and Weber (1958) reported from Mt. McKinley Park sterile but "well developed" material of an undeterminable species of Fossombronia. (It is entered in the present list as "Fossombronia sp.") Finally, all collections from British Columbia previously determined as Marsupella ustulata are not that species according to R. Grolle (pers. comm.). It is possible that the same is true for the Alaskan material.