Arctoa (2015) 24: 327-361 doi: 10.15298/arctoa.24.28 LIVERWORTS FROM THE LARGEST OF THE UNITED STATES: A CHECKLIST FOR MAINLAND ALASKA ЧЕКЛИСТ ПЕЧЕНОЧНИКОВ КОНТИНЕНТАЛЬНОЙ АЛЯСКИ, КРУПНЕЙШЕГО ШТАТА США LARS SÖDERSTRÖM1, ANDERS HAGBORG2 & MATT J. VON KONRAT2 ЛАРС СЁДЕРСТРЁМ1, АНДЕРС ХАГБОРГ2, МАТТ Д. ФОН КОНРАТ2 Abstract We report 294 liverwort taxa in 267 species for the State of Alaska, excluding the Aleutian Islands,15 taxa are listed as doubtful and an additional 17 of the previous reports are rejected as erroneous. The checklist is based on 417 literature references, including monographs, regional studies, and molecular investigations. The list represents the first compilation of liverworts from the region for over three de- cades. It is likely that a number of new records remain to be reported, based on regionally widespread species that have been recorded elsewhere. On the other hand, it is envisaged that a number of the taxa presented here may be synonyms of other taxa as they are taxonomically insufficiently known. Резюме Чеклист включает 294 таксонов печеночников, относящихся к 267 видам, известным с Аляски, не включая Алеутские острова. 15 таксонов отнесены к сомнительным, а eщe 17 исключены из флоры, поскольку выяснилась ошибочность определений. Чеклист основывается на 417 литературных источниках, в том числе систематических обработках, региональных флорах и заметках, а также образцах, цитированных в молекулярно-филогенетических работах. Список представляет собой первое за последние 30 лет обобщение по региону. Новые находки здесь вполне вероятны, учитывая, что не все распространенные на сопредельных территориях виды уже найдены в штате. С другой стороны, нельзя исключать что дальнейшие таксономические исследования приведут к сведению части таксонов в синонимы. KEYWORDS: Alaska, liverworts, checklist. INTRODUCTION scapes, the bryophyte flora remains relatively poorly Bryophytes are a major component of Alaska’s eco- known in comparison to the vascular flora and wildlife systems, as in many northern landscapes, dominating generally. This is especially the case for liverworts. plant biomass and production (Wielgolaski et al., 1981). We here present a checklist of liverworts for the State Bryophytes, together with other cryptogams generally, of Alaska (excluding the Aleutians) to summarize the play critical roles in the functioning of many northern current knowledge and further promote and encourage ecosystems of high latitude (Longton, 1997; Cornelissen bryological research in the region. The significance of et al., 2007) such as regulation of hydrology (Beringer et checklists was summarized by Söderström et al. (2007, al., 2001), carbon balance (Rydin & Jeglum, 2006), ni- 2008), outlining the utility of checklists as powerful and trogen fixation (Solheim et al., 1996), soil drainage (Vitt important tools, and their applicability to taxonomy, sys- & Pakarinen, 1977) and preservation of permafrost (Dyr- tematics, and conservation. ness, 1982; Yoshikawa et al., 2002). Given their impor- As with many other regions in the world, given the tance, changes in abundance and species composition of relatively poor historical focus on liverworts in Alaska, cryptogams in northern landscapes have significant im- it is likely that a vast number of new records are yet to be pacts on regional biogeochemistry and feedbacks to glo- reported. In this checklist we report 294 taxa in 267 spe- bal climate (Lang et al., 2011). Around the world, in- cies, with another 14 taxa questioned. Seventeen taxa cluding Alaska, oil and gas exploration and development are rejected as erroneously reported from Alaska. are moving into areas previously undisturbed by indus- Alaska is the largest of the United States. The conti- trial development (Wilson et al., 2013). Yet, despite the nental part covers 16 degrees of latitude stretching from ecological significance of bryophytes in northern land- Lake and Peninsula Borough at 55°N to Point Barrow at 1 – Department of Biology, Norwegian University of Science and Technology, N-7498 Trondheim, Norway. E-mail: [email protected] 2 – Department of Science and Education, The Gantz Family Collections Center, The Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605, U.S.A. Email: [email protected]; [email protected] 328 L. SÖDERSTRÖM, A. HAGBORG & M.J. VON KONRAT Fig. 1. Terrestrial ecoregions of Alaska following Olson et al. (2001). 71°N. It covers 38 degrees of longitude from Ketchikan such as Alaska and elsewhere in the Arctic, is the vast- Gateway at 130°W to Nome census area at 168°W. Alas- ness and the inaccessibility of many or most regions ka is ecologically diverse ranging from coastal to upland (Gillespie et al., 2012). Despite this, there have been tundra, coastal to boreal forests, to vast ice fields (Fig. many avid collectors and an active interest in liverworts, 1). Wahrhaftig (1965) reported the great topographic di- especially since the 1950’s. Interestingly, Alaska is the versity of Alaska in detail and divided it into 12 physio- type locality for 23 described taxa, eight of which are graphic provinces and 60 smaller divisions. recognized by current taxonomy. It remains very evi- Hultén (1940) provided a detailed history of early dent that Alaska, in general, remains under-explored botanical exploration in the Alaska and Yukon territo- for liverworts and much yet is to be discovered. It is ries from the time of their discovery by Europeans in interesting that no hornworts seem to have been re- 1741 to the year 1940. Hultén provided a comprehen- ported from Alaska, or any other area of the American sive list of early collectors, including details of their arctic. exploration and voyages, collections, herbaria housed, MATERIALS AND METHODS and resulting publications. Steere (1978), Steere & In- Nomenclature and taxonomy follows the forthcom- oue (1978) and Davison (1993) provided historical ac- ing world checklist of hornworts and liverworts (Söder- counts of bryology in Alaska, including the North Amer- ström et al. 2015). Sources include 417 publications found ican Arctic in general. Steere (1978), focusing on the through the work with Early Land Plants Today (ELPT) Arctic, remarked that the bryophytes of Arctic Alaska database of liverwort taxonomy and distribution, and with remained almost totally unknown until the 1950’s, in some consultation with taxonomic experts. The check- stark contrast to Siberia, Arctic Canada and Greenland list follows a similar format of previous liverwort and that were relatively well known. The historically poor hornwort checklists by the authors, e.g., Java (Söderström knowledge of liverworts from Alaska had its roots in et al., 2010), Fiji (Söderström et al., 2011a), and Jamai- 18th and 19th century explorations and is summarized ca (Söderström et al., 2011b). The checklist distinguish- by an example offered by Steere & Inoue (1978). They es between records that are based on specimens (refer- noted that despite significant collections of plants, in- ence in bold) and second-hand reports, e.g., citations of cluding bryophytes, made by several important voyages earlier publications (reference in normal type). All names of exploration in the first half of the 19th century relat- used for the references are given in brackets after the ing to a search for a Northwest Passage through the taxon name with spelling variants/errors within quota- seas north of North America, only one species of liver- tion marks. Taxa are arranged in alphabetic order. Sig- wort was collected and recorded – the cosmopolitan nificantly, each accepted taxon is qualified using a four Marchantia polymorpha. One major challenge in areas level ranking system that indicates our level of knowl- Liverworts from the largest of the United States: a checklist for mainland Alaska 329 Fig. 2. Geographical regions used and the num- ber of species occurring in each region. For the ab- breviation of the regions, see text. The darker brown, the more species are reported from the region. The abbreviations are given in Table 1. Table 1. Abbreviations of regions of Alaska (see also Fig. 2) ACCEPTED TAXA AN Anchorage Borough Acrobolbus *** A. ciliatus (Mitt.) Schiffn. ASK: Worley 1970, Hattori BB Bristol Bay Borough 1975, Sharp 1987, Long & Grolle 1990. PH: Schofield 1988. BE Bethel census area DE Denali Borough Anastrepta DI Dillingham census area *** A. orcadensis (Hook.) Schiffn. ASK: Brinkman 1934, Frye FB Fairbanks North Star Borough & Clark 1945, Müller 1954b, Grolle 1966b, Hattori 1966, 1968, 1971, 1975, Schofield 1968, Worley 1970, Schofield KE Kenai Peninsula & Crum 1972, Damsholt 2002. KI: Persson 1946. PH: Evans KI Kodiak Island Borough 1915b, Persson 1946, Worley 1972. VC: Persson 1946, Ey- LP Lake and Peninsula Borough erdam 1952, Worley 1972. MS Matanuska-Susitna Borough Anastrophyllum NO Nome census area *** A. assimile (Mitt.) Steph. (Anastrophyllum reichardtii “re- NS North Slope Borough ichardti”). ASK: Stephani 1901c, Evans 1915b, Herzog NW Northwest Arctic Borough 1926, Arnell 1928, Brinkman 1934, Evans & Nichols 1935, PH Panhandle – this is a combination of ten Frye & Clark 1945, Arnell 1950, 1956, Müller 1954b, Grolle boroughs and census areas in the southeast of 1964b, 1966b, Hattori 1966, 1968, 1975, Hong 1966, 1996, Alaska Worley 1970, Schuster & Damsholt 1974, Godfrey 1977b, SF Southeast Fairbanks census area Long & Grolle 1990, Váňa 1991, Bapna & Kachroo 2000b, VC Valdez-Cordova census area Konstantinova 2000, Damsholt 2002, 2013, Schill & Long WH Wade