The sporophyte of Bucklandiella affinis under magnification with Mt. Johannesburg in the distance.
2019 North Cascades National Park Bryophyte Inventory Project and the Sub-glacial Bryophyte Survey
Summary
The North Cascades National Park (NOCA), located near the Pacific coast of western North America centre of endemism – as specified in the IUCN SSC Bryophyte Action Plan (Hallingbäck & Hodgetts 2000) – hosts populations of bryophyte genera both globally and regionally rare. Despite its presence in one of the largest contiguous wilderness areas in the United States, with over 550,000 square acres of deep river basins and heavily glaciated mountain ranges, it has relatively sparse bryophyte records. Since 2016 I have worked to expand the known bryophyte taxa from the Park, as well as the adjacent forest service lands. My NPS endorsed bryophyte collection trips have largely focused on subalpine and alpine outcroppings, seepages and wetlands, as these areas are the least collected from in the park. In the past three years of bryophyte surveys in the North Cascades Range, I have added 225 bryophyte specimens to the park’s herbarium, twenty of which are common and rare species newly documented for the Park.
Bryophyte inventorying prospects for 2019 in NOCA will continue to prioritize collection trips to subalpine and alpine zones. Areas of interest are the high elevation talus fields and outcroppings of the northwestern part of the Park where the major rock type is Shuksan Greenschist. This rock is partially composed of calcium-aluminum (Lawsonite) thereby giving it a high pH signature. Collections will thus focus on bryophytes of high elevation (900m – 2100m) wetlands, snowfield seepages, and outcroppings with the anticipation of finding calciphilous bryophyte species as well as the 25 rare mosses and liverworts listed in Tables 1 & 2. Due to the rarity of alkaline rock in the Pacific Northwest, especially at high elevations, any collections of calciphilous species would be significant. This study area will also be suitable for surveying for Brachydontium olympicum, B. trichoides, Bryoxiphium norvegicum subsp. norvegicum, Bartramiopsis lescurii and Grimmia lesherae, as these globally threatened and imperiled species have adjacently known populations. Since this region of the park is entirely lacking in bryophyte records, any collections from this area will provide valuable information for the inventorying and monitoring efforts of the North Cascades National Park, and bryophyte conservation efforts at large.
In addition to the previously mentioned bryophyte collection goals, we will be surveying the margins of alpine glaciers in the vicinity of the Berdeen unit for Little Ice Age (LIA) entombed bryophyte remains. Three glaciers in the North Cascades Range were initially surveyed for such remains in 2018 but were not found. Myself and those that have helped are essentially asking the question, “can alpine bryophyte communities endure glacial entombment and, once exhumed, continue biological activity via totipotency (regrowth from stem and lateral branch apices)?”
Bryophytes to be surveyed for of highest conservation concern:
Bartramiopsis lescurii (James) Kindberg
- Conservation status: State: S1 (Critically imperiled); Global status: G3G5 (Imperiled-Secure)
- Known distribution: B.C., Alaska, Washington, East Asia
- Locations within Washington State: 1 location in Cascade Mountains.
- Habitat association: Soil banks or soil over rock, exposed tree roots; usually low to moderate elevations (eFloras, 2019)
Brachydontium olympicum (E. Britton) T. T. McIntosh & J. R. Spence
– Conservation status: State: S1 (Critically Imperiled); Global status: G2G3 (Imperiled-Vulnerable)
– Known distribution: Japan, Alaska, British Columbia, Washington, and Oregon
– Locations within Washington state: 3 locations in Mt. Rainier, Jefferson County, and the
North Cascades Range.
– Habitat Association: Growing on exposed boulders, or thin soil in rock clefts. Mostly
found above timberline between 5,000- 6,000 ft. (Christy,2007)
Brachydontium trichoides (F. Weber) Fürnrohr
– Conservation status: State: S1 (critically imperiled); Global status: G3 (Vulnerable)
– Known distribution: Northern and central Europe, China, Columbia, Australia
Tennessee, North Carolina, New Hampshire, and Mt. Rainier, Washington.
(eFloras, 2019)
– Locations within Washington State: One location within Mt. Rainier National Park.
– Habitat Association: saxicolous – sand, or rock, overhung and vertical surfaces.
Bryoxiphium norvegicum subsp. norvegicum (Bridel) Mitten, J. Linn.
– Conservation status: State: S1 (Critically imperiled); Global status: G3 (Vulnerable)
– Known distribution: Europe, North America: Alabama, Alaska, Arizona, Arkansas,
Indiana, Iowa, Kentucky, Minnesota, Missouri, North Carolina, New Mexico, Ohio,
Tennessee, Washington, and Wisconsin (eFloras, 2019)
– Locations within Washington state: 1 location in the Olympic mountains, 1 location in
Snohomish county, Mt. Rainier.
– Habitat association: Growing on porous rock ledges and faces, sometimes overhanging
water. Calcareous tendencies have been reported, but more often volcanic, sandstone,
or gneiss substrates. (eFloras, 2019)
Grimmia lesherae Greven
- Conservation status: State: not assigned; Global status: of conservation concern
-Known distribution: Washington, California
-Locations within Washington: one location, very close to study site in the North Cascades Range.
- Habitat association: Damp acidic rock; of conservation concern; high elevations (2000-3000 m).
Hookeria acutifolia Hooker & Greville Conservation Status: S1; G4G5
Meesia triquetra (Linnaeus ex Jolyclerc) Ångström Conservation Status: S1; G5
Meesia uliginosa Hedwig Conservation Status: S1; G4
Myurella julacea (Schwagrichen) Schimper Conservation Status: S1; G5
Neckera pennata Hedwig Conservation Status: S1; G5
Orthothecium chryseum (Schwägrichen) Schimper var. chryseum Conservation status: S1; G5?
Orthotrichum hallii Sullivant & Lesquereux Conservation status: S1; G4
Orthotrichum pylaisii Bridel Conservation status: S1; G4G5
Pohlia cardotii (Renauld) Brotherus Conservation status: S1; G2G3
Rhytidium rugosum (Hedwig) Kindberg Conservation Status: S1; G5
Schistostega pennata (Hedwig) F. Weber & D. Mohr Conservation Status: S2; G3G4
Tetradontium brownianum (Dickson) Schwägrichen var. brownianum Conservation Status: S1; G3G4
Tomentypnum nitens Hedwig Conservation Status: SNR; G5
Table 1. Rare mosses[1] to be surveyed for. Conservation status is provided in Washington State ranking (S) and global ranking (G). S(1) – Critically Imperiled S(2) -Imperiled G(2) – Imperiled G(3) – Vulnerable G(4)- Apparently Secure G(5)- Secure SNR - Unreviewed
Gymnomitrion concinnatum (Lightf.) Corda
Haplomitrium hookeri (Lyell ex. Sm.)
Harpanthus flotovianus (Nees) Nees
Herbertus aduncus (Dicks.) Gray
Herbertus sakuraii (Warnst.) S. Hatt.
Lophozia laxa (Lindb.) Groelle
Marsupella condensata (Ångstr. ex C. Hartm.) Lindb. ex Kaal.
Marsupella sparsifolia (Lindb.) Dumort.
Nardia japonica Steph.
Radula Brunnea Steph.
Scapania gymnostomophila Kaal.
Scapania obscura (Arnell & C.E.O. Jensen) Schiffn.
Table 2. Rare liverworts[2] to be surveyed for
Figure 1. Showing the boundaries of the North Cascades National Park Complex, 2019 bryophyte inventory site, and the Rainy Pass bryophyte inventory site from 2016.
NOCA Bryophyte Inventory Project Background
The North Cascades Range, located in the Northwestern part of the state of Washington, is an assemblage of geological terrains, diverse climates, and biological communities. A phytogeographic interest towards bryophytes in the North Cascades has been growing since the early 1980's when John Spence reported disjunct populations of dry, interior Rocky mountain bryophyte taxa on Chowder Ridge (Spence, 1986), and about 12 km east, Wilfred Schofield found several populations of the globally imperiled Brachydontium olympicum in the early fall of 1981. Eighteen years later and about 10 miles south east, Robin Lesher - a botanist with the USFS - was conducting plant inventories in a talus field on the side of Snowking Mountain when she stumbled upon a population of Grimmia with an uncharacteristically straight seta. This specimen went through tedious species determination efforts until it was determined to be a distinct species, Grimmia lesherae, and has since only been found in one other location: near Mt. Shasta.
Bryophyte inventories in the North Cascades since the time of Spence and Lesher have largely been focused in the western periphery of the range while the heavily glaciated interior of the range – the North Cascades National Park – still has exceptionally sparse bryophyte records[3]. The over 550,000 acres of the North Cascades National Park have been visited by trained bryologists only a handful of times. Most noteworthy bryophyte collections have been by Dan Norris when in September of 1999 he collected 156 species of mosses and liverworts from 1000 – 2000m in the eastern and southern part of the Park. In 2004 Martin Hutten contributed an immense collection of bryophyte and liverwort taxa from the Skagit Gorge[4], a very narrow 4-mile stretch of the Skagit river. His taxonomic index provided a thorough knowledge of bryophyte taxa from the lower elevations of the adjacent Ross Lake National Recreation Area. However, most of the park remains largely unexplored by bryologists.
In 2016 I began conducting bryophyte inventories within the North Cascades National Park and the adjacent Okanogan-Wenatchee National Forest. In 2016 the International Association of Bryologists generously provided me with funding to survey for Brachydontium olympicum, Brachydontium trichoides, and Bryoxiphium norvegicum subsp. Norvegicum within the Rainy Pass area, just east of the Park boundary. This project resulted in the documentation of 77 common and rare[5] moss and liverwort taxa (although the three original species of interest were not found.)
My bryophyte collection efforts in NOCA have so far focused on inventorying from wetlands and subalpine and alpine sites. Most noteworthy among the rare collections have been Andreaea rothii[6] and tetraplodon mnioides.[7] The newly introduced documentation of six common acidic wetland obligate species of Sphagnum[8] infers a lack of acidic wetland bryophyte surveys at large for the Park.
Subglacial Bryophyte Survey
Since the late 1800's, approximately 56% of the North Cascades National Park's total ice cover has been lost (Riedel & Larrabee, 2018) reaching the greatest mass minimum in the past 4,000 years (Osborn et. al. 2012). As glacial ice continues to retreat to higher elevations, exposed moraines, lakes, and other topographic features have produced plant materials which have been used to date glacial advances and retreats during the Holocene (Osborn et. al., 2012) and infer likely vegetative communities and locations of biological refugia at that time. Established biological refugia concepts for mountain species identify three types of refugia: peripheral (ridgelines between glaciers), lowland (below glaciers), and nunatak (above glaciers) (Holderegger & Thiel-Egenter, 2009). However, recent research has established a fourth biological refugia scenario that might be possible in the North Cascades Range.
In 2007 Dr. Catherine LaFarge and colleagues collected intact bryophyte specimens immediately adjacent to the terminus of the receding Teardrop Glacier on Ellesmere Island in the Canadian Arctic. Upon receiving the radio carbon dates of the specimens, it was determined that they had found remnants of the last inter-glaciation which had endured glacial entombment through the Little Ice Age. In vitro growth experiments showed that some specimens regenerated - after nearly 450 years of entombment - via stem apices and lateral branches, thus demonstrating totipotent behavior in response to exposure (LaFarge et. al., 2013). The team’s findings help further the discussion on ice age refugia concepts of tundra communities, and possibly temperate alpine colonization as well. If glacially entombed bryophytes from past inter-glaciations play a role in re-colonization of alpine terrain in the North Cascades, this may have significant implications for understanding the evolution of western North American alpine bryophyte taxa and would allow theories of biological refugia in the Cascades Range to be revisited.
In conversation with Dr. LaFarge about her work on this phenomenon, I inquired if LIA bryophyte re-emergence from alpine glaciers in temperate regions has been studied. To her knowledge there has been no such research, and surveying for similar findings as in the Teardrop Glacier would be worthwhile. In 2018, with advice from Dan Smith (alpine geomorphologist and dendrochronologist, with the University of Victoria) and John Riedel (glaciologist and geologist from the North Cascades National Park), I selected glaciers in the North Cascades that fit their suggested parameters for finding LIA bryophyte remains. The results from these surveys did not yield any such remains that LaFarge found in her study sites but did serve to further develop the scientific protocol of this study.
The ‘dying’ Colonial Glacier.
An old USGS gauging station was placed many years ago at the discharge point of the lake at the base of the Little Devil Glacier.
Eve Bernhard and I encountered a late summer snowfall which made searching for LIA bryophytes at the Quien Sabe Glacier, needless to say, quite difficult.
Activities:
The Subglacial Bryophyte Survey and the Bryophyte Inventory project will be conducted during the glacial minimum mass balance which will be from August through September of 2019 in the Berdeen unit of the North Cascades National Park. This is a 90 sq. km area characterized by unique geology, glaciated topography and high elevation wetland basins where no bryophyte records exist. During this time window there will be at least one bryophyte collection trip consisting of three trained bryologists who will intensively survey talus fields, snowfield seepages, outcroppings and other sites that might provide suitable habitat for Brachydontium olympicum, Bryoxiphium norvegicum subsp. norvegicum, Bartramiopsis lescurii and Grimmia lesherae along with the listed rare mosses and liverworts in tables 1 and 2. Each specimen collected will be returned to the lab in Marblemount and identified using the Park’s provided resources. Specimens will then be vouchered with retrievable data such as GPS (UTM), qualitative description of the site (habitat, associated vegetation, aspect, and substrate), sporophyte presence, population size (extent in area units). In addition, we plan to survey the margins of the Noisy Glacier on the north aspect of Bacon Peak, and other low gradient glaciers on its south and east flanks during the bryophyte collection trips for LIA bryophyte remains. If we do find LIA bryophyte remains, the specimens will be placed in the freezer in the North Cascades National Park’s research station. The scope and budget of this project only covers the preliminary survey work for LIA bryophyte remains. If such specimens are collected, they will be frozen until further research can be conducted.
Rationale
In 2016 the IAB generously provided me funding to survey for these species in the Rainy Pass area of the North Cascades. My work produced a list of 77 rare and common bryophytes for the area, although I did not locate the target species. After three years of rigorous bryophyte inventorying in the North Cascades, I have identified this area of the park (Berdeen unit) to possibly support communities of rare bryophyte taxa. The unique geology, glaciated topography and adjacency to one of the only known locations of several of these globally imperiled species makes this location paramount for furthering the development of alpine bryophyte floristics and possibly broadening our understanding of temperate alpine ice age refugia. Furthermore, high elevation bryophyte surveys such as this project are essentially important considering the correlation between alpine zone recession and climate change. Current climate change projections for sparsely vegetated uplands in the North Pacific region anticipate advancements in tree-line (Rocchio, and Crawford, 2015) and a potential loss of habitat for alpine specific species. Considering this risk, and the small population size and geographic continuity of these species, this places them in threat of regional extirpation.
Expected outputs
Upon completion of this study, all collected vouchered specimens will be given to NOCA’s herbarium. A report of our findings will also be produced and given to the Resources division of NOCA, the Washington State Department of Natural Resources’ Natural Heritage Program for species conservation status review, the International Association of Bryologists and other interested agencies and colleagues. Depending on the amount of new information produced from this study, a report will be provided to the IUCN and an article may be submitted to the Northwest Science Association for publication.
References
Christy John. A. 2007. Species fact sheet: Brachydontium olympicum. US Forest service, 2009
'eFloras (2019). Published on the Internet http://www.efloras.org [accessed 22 April 2019]' Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
Hallingbäck, T. and Hodgetts, N. (compilers). (2000). Mosses, Liverworts, and Hornworts. Status, Survey and Conservation Action Plan for Bryophytes. IUCN/SSC Bryophyte Specialist Group. IUCN, Gland, Switzerland and Cambridge, UK. x + 106pp.
Harpel, Judy et. al. 2015 Washington Natural Heritage Program List of Mosses. Washington Department of Natural Resources.Holderegger, R. and Thiel‐Egenter, C. (2009), A discussion of different types of glacial refugia used in mountain biogeography and phylogeography. Journal of Biogeography, 36: 476-480. doi:10.1111/j.1365-2699.2008.02027.xOsborn et. al., 2012. Latest Pleistocene and Holocene glacier Fluctuations on Mt. Baker, Washington. Quaternary Science Reviews. 49 (2012) 33-51Regeneration of exhumed Little Ice Age bryophytes. Catherine La Farge, Krista H. Williams, John H. England. Proceedings of the National Academy of Sciences Jun 2013, 110 (24) 9839-9844; DOI:10.1073/pnas.1304199110
Riedel J and Larrabee MA. 2018. North Cascades National Park Complex glacier mass balance monitoring annual report, water year 2013: North Coast and Cascades Network. Natural Resource Data Series. NPS/NCCN/NRDS—2018/1142. National Park Service. Fort Collins, Colorado
Rocchio, F. Joseph, Crawford, Rex. 2015 Ecological Systems of Washington State. A Guide to Identification. Natural Heritage Report 2015-04. Washington Department of Natural Resources, Natural Heritage Program. Pp: 2012-2015.
Spence, J. R. 1986. The moss flora of the alpine – high subalpine Chowder Ridge area, western North Cascades Range, Washington State, U.S.A. Can. J. Bot 64: 146-150.
Tropicos.org. Missouri Botanical Garden. 30 Jun 2017 <http://www.tropicos.org>
© 2017 Missouri Botanical Garden - 4344 Shaw Boulevard - Saint Louis, Missouri 63110
[1] Conservation statuses adopted from Harpel et. al., 2015
[2]Authorities in Tropicos Taxanomic nomenclature
[3] NOCA’s herbarium has a total of 1200 archived non-vascular specimens consisting of 253 species of mosses, 86 species of liverworts, and 0 hornworts; Compare to Olympic NP which has over 13,000 specimens.
[4] The Skagit River gorge as well the three reservoirs created by Seattle City Light hydroelectric power company are part of the Ross Lake National Recreation Area. The North Cascades National Park surrounds this area and are managed by the Park, but under different regulations.
[5] Anastrophyllum sphenoloboides, Anastrophyllum minutum, Marsupella sparsifolia, Nardia breidleri
[6] S1G5 Conservation Status.
[7] S1G5 Conservation Status.
[8] Sphagnum compactum, S. fimbriatum ssp. fimbriatum, S. miyabeanum, S. palustre, S. rubellum, & S. subnitens
Bucklandiella affinis and the Mt. Johannesburg massif in the background.
2016 - 2017 Bryophtye Surveys in the North Cascades National Park Complex
Subalpine and alpine zones, montane riparian sites and acidic wetlands were the focus for bryophyte collection field trips within the NCNPC during 2016, 2017 and 2018 field seasons. Combined, 100 liverwort and moss species were documented throughout the collection sites. Rare liverwort species collected from NCNPC were: Radula obtusiloba subsp. polyclada (A. Evans) S. Hatt., and Asterella lindenbergiana (Corda ex Nees) Lindb. ex Arnell. One of the mosses collected were shown to have a conservation status of concern: Andreaea rothii F. Weber & D. Mohr. Six Sphagnum species and three other wetland obligate species were newly documented for the Park. Combined, 20 bryophyte species were added to the park’s inventory. Such developments for NCNPC’s bryophyte inventory indicate its acidic wetlands to have been poorly surveyed for bryophytes thus far, and likely support more undocumented taxa. The bulk of the species collected were common to the North Cascades.
Bryophyte taxa newly added to the Park’s inventory:
Andrea rothii F. Weber & D. Mohr
Asterella lindenbergiana (Corda) Lindb.
Calliergon giganteum (Schimper) Kindberg
Dichelyma urnicatum Mitten
Dicranoweisia crispula (Hedwig) Milde
Fissidens dubius P. Beauvois, Prodr.
Jungermania exertifolia Steph. Ssp. cordifolia (Dum.) Vana
Jungermannia obovata Nees
Leicolea heterocolpos (Thed.) Bruch
Pogonatum dentatum (Bridel) Bridel
Pohlia wahlenbergii (Web & Mohr.)
Polytrichastrum alpinum var. Septentrionale Brid.
Racomitrium obesum Frisvoll
Schistidium duprettii (Theriot) W.A. Weber
Sphagnum compactum Lam. & DC
Sphagnum fimbriatum ssp. fimbriatum Wilson & Hooker
Sphagnum miyabeanum Warnstorf, Carl (Friedrich E.)
Sphagnum palustre Linneaus
Sphagnum rubellum Wilson
Sphagnum subnitens Russow & Warnst
Bryophyte Inventories: 2016
A survey for rare bryophytes: Brachydontium olympicum, Brachydontium trichodes, and Bryoxiphium norvegicum in the northern Cascade Range of Washington State.
This work started in June of 2016 with the generous Conservation and Endangered Species Grant from the International Association of Bryologists and the University of Connecticut. This survey sought primarily to locate and describe possible locations of the above species, but also to develop and inventory the sparsely documented bryophyte taxa from the Rainy pass area.
Abstract
Bryophyte collection trips from Rainy Pass, within the Okanogan-Wenatchee National Forest, largely focused on surveying for the globally imperiled mosses – Brachydontium trichodes (F. Weber) Milde, Brachydontium olympicum (E. Britton) T.T. McIntosh & J.R. Spence, and Bryoxiphium norvegicum subsp. norvegicum (Brid.) Á. Löve and D. Löve – among the subalpine and alpine zones of Rainy Pass. Both the accessibility, and high elevation, in proximity to known locations for these species allowed for Rainy Pass to be a suitable area to survey. Due to the sparse bryophyte records from the North Cascades, all species encountered were collected such that a comprehensive bryophyte species list could be developed for the area. 77 species were collected from throughout the pass. Three liverworts from the collections show to have few records from Washington. The three species of interest were not found from the Rainy Pass study area.