MONTANA'S ARCTIC FLORA

The first time I came to Montana I went hiking above tree line in Glacier National Park. The views were spectacular, but just as exciting to me were the plants, many of which had "arctic-alpine" distributions -- plants of the frozen north right here in Montana. Over 100 of the nearly 2,100 native plants in Montana (ca. 5%) belong to the Arctic Flora, a group of species that has evolved under an arctic climate and soils over long periods of time. The Arctic Flora dominates the vegetation north of the Arctic Circle (ca. 67º N latitude). A strict botanical definition of arctic pertains to those lands north of the limit of well formed trees (i.e., full crown, straight, erect bole). In Canada this includes Baffin Island, all of the area around Hudson's Bay, as well as that north of Great Slave Lake. It also includes the northern one-third of Alaska. Plant species belonging to the Arctic Flora are obviously adapted to a cold, often cloudy climate. However, the growing season, during which the upper layers of soil have thawed and temperature is above freezing during the day, can be surprisingly long compared to many alpine environments. North American arctic soils are either derived from acidic Precambrian bedrock (Canadian Shield) or from calcareous Paleozoic, sedimentary rock.

North American arctic vegetation has been classified into many types, but here I will use a simple, dichotomous classification: (1) polar desert/fellfield with well-drained, poorly-developed soil and (2) tundra with relatively poor drainage (often due to permafrost) and well-developed, moist to wet soils. Arctic plants occur in similar habitats above treeline in alpine settings south of the arctic.

Nearly half of Montana's 108 arctic plants occur in only four families: Cyperaceae (15), Asteraceae (12), Saxifragaceae (12), and Rosaceae (9). Three of our arctic species are annuals, three are dwarf shrubs, five are shrubs, and the rest are herbaceous perennials. Thirty-four of the 108 arctic-alpine species that occur in Montana are considered rare here because they are on the southern margin of their range or have a disjunct or fragmented distribution in the state. The majority (71%) of these rare- in-Montana arctic plants occur in only one or both of two areas of the state: (1) Glacier National Park and the adjacent Flathead and Front ranges in the northwest and (2) the Beartooth Range and nearby Crazy Mountains in south-central Montana.

The Lewis Range of Glacier National Park receives more snow than anywhere else in Montana due to the prevailing Jet Stream. As a result it can support wet tundra, although this habitat is limited due to steep topography and deep snow. The geology of the Glacier-Front region also provides limestone fellfields. These habitats with poorly-developed soil apparently simulate the polar desert/ fellfield habitats of the arctic. Rare arctic species found in this habitat include Saussurea nuda, Carex petricosa, Carex glacialis, Arnica angustifolia, Braya humilis, Dryas integrifolia, and Senecio cymbalaria. The former three are confined to the Glacier-Front region, while the latter four occur elsewhere. Although most of these species are associated with both acidic and calcareous parent materials in the Arctic, they are confined to limestone-derived soils in Montana.

William Weber, who has studied the arctic-alpine flora of Colorado extensively, pointed out that many arctic disjuncts occur in east-west trending mountain ranges.

The north slopes of these exceptional ranges (most Rocky Mountain ranges trend north-south) are thought to have acted as refugia from the exceptionally warm climate approximately 8,000 years ago during the altithermal period. This pattern holds true for Montana as well, because the east-west-trending Beartooth Range is the premier refugium for arctic disjunct species in the state; 16 of our rare arctic species are found there, and all but one of these are found in seeps or moist to wet tundra .

A more restrictive habitat association I have noticed is that many of our rare arctic tundra species are found not on north-facing slopes but rather on gentle, south- or west-facing slopes below permanent snow fields. Such species include Draba macounii, Eriophorum callitrix, Kobresia simpliciuscula, Kobresia sibirica, Ranunculus sulphurous, and Tofieldia pusilla. It is my hypothesis that the south exposure provides a long growing season, and the meltwater from permanent snow assures that the soil remains wet and cold, similar to arctic tundra during the summer.

The presence of arctic species in Montana suggests that the arctic flora "migrated" down the Rocky Mountain chain during the cooler Pleistocene epoch. These plants are a window on the past and may provide clues to how our vegetation may respond to future climate changes. Nearly one-quarter of our 34 rare arctic species was first discovered in Montana in the past 25 years. There are undoubtedly more to be found, so keep your eyes open.

Table - Habitat and distribution of rare arctic species in Montana

References

Aiken, S.G., Dallwitz, M.J., Consaul, L.L., McJannet, C.L., Boles, R.L., Argus, G.W., Gillett, J.M., Scott, P.J., Elven, R., LeBlanc, M.C., Gillespie, L.J., Brysting, A.K., Solstad, H., and Harris, J.G. 2007.

Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. NRC Research Press, National Research Council of Canada, Ottawa. http://nature.ca/aaflora/data

Panarctic Flora Project Steering Committee. Undated.

Annotated Checklist of the Panarctic Flora (PAF) Vascular plants. http://nhm2.uio.no/paf/

Weber, W. A. 2003.

The Middle Asian element in the Southern Rocky Mountain flora of the western United States: a critical biogeographical review. Journal of Biogeography 30: 649-685.

FITTING IT IN: MAKING ROOM FOR THE SEDGES OF THE PACIFIC NORTHWEST--ALL OF THEM

Field Guide to the Sedges of the Pacific Northwest - Second Edition by Barbara L. Wilson, Richard Brainerd, Danna Lytjen, Bruce Newhouse and Nick Otting The Carex Working Group Published by the Oregon State University Press, Corvallis, OR. ISBN 978-0- 87071-728-4 [soft cover] 432 p. Price $35.00

Order from OSU Press http://osupress.oregonstate.edu/book/field-guide-to-sedges-of-pacific-northwest-0

Additional information available from http://www.carexworkinggroup.com/pages/fieldguide.html

We, Carex Working Group members, were excited that OSU Press agreed to publish a second edition of our Field Guide to the Sedges of the Pacific Northwest. This would be an opportunity to revise the identification key, update nomenclature, modify a few maps, improve some of the photos, and make minor updates and corrections in the text. And clarify the Carex subbracteata confusion. Definitely clarify that.

The process became more exciting than we intended.

Our first big concern was adding Carex species recently discovered in Oregon or Washington. Each species needed a page of text plus a page of pictures. We needed to keep the number of pages constant. Additions included natives Carex eburnea from northeast Washington and C. subbracteata from the coast, as well as introduced C. distans, C. divulsa, and C. hirta. (Fuzzy Carex hirta was collected in Portland a century ago, rediscovered in 2010.)

We decided to go with the split in Carex capillaris, which meant adding native Carex tiogana from Steens Mountain. Let's see . . . six added sedges meant twelve added pages. We saved two pages by recognizing only Carex heteroneura, not its two subspecies that couldn't really be distinguished. That left ten pages to find. Where?

We cut "Excluded, Extirpated, and Not (Yet?) Discovered Species" from five pages to two since most of the species covered there had been found or never would be. Reformatting "Sedges with Distinctive Traits or Habitats" gained a couple more pages. Using a smaller font in the index helped, too. We sacrificed the ethnobotany section; much though we liked it, it didn't help people identify sedges and we needed its four pages. (We tucked as much ethnobotany into individual species accounts as we could.)

Updating the key was simple because we had been revising our personal version of it all along. Improving the text was fun. Finding photos of the new sedges was harder, especially since we wanted to be sure the identifications were correct. We ended up corresponding with botanists from Michigan, Illinois, Germany, and Romania to get the photos we needed.

Sorting out Carex subbracteata and C. harfordii took a lot of correspondence and much looking at specimens, but we did it. We hope. Both species have full treatments now.

In late summer 2013, as we neared the end of our rewriting, we learned that Chris Reidy and Kathy Pendergrass from the Natural Resources Conservation Service had found Carex hirsutella, native to eastern North America, on a farm in Linn County. We went out to investigate and confirmed the report. This was a very interesting find, but our main concern was "Two more pages! We need two more pages!" Fortunately, we discovered that we had misidentified Carex projecta. Removing its account freed up the needed pages.

At last the revision was completed. We took a jump drive with the text and photos to OSU Press. There, our wonderful editor Jo Alexander told us exactly what she thought of the hundreds of changes we had made. This was supposed to be a revision, not a new book! Yes, of course, she would make the changes, but we were asking a lot! Finally we got proofs and carefully read through it again, of course finding small errors we had missed.

Then Rex Crawford and Joe Rocchio of the Washington Department of Natural Resources reported that they had found Carex lacustris in far northeast Washington. Aaaargh!!!!! We had heard that Carex lacustris probably grew in northeast Washington but there were no specimens to confirm its presence there. We'd mentioned it in the first edition, even put it in the key, but since no one found it we omitted it from the second edition to save space.

When photos confirmed that this new report of Carex lacustris was accurate, we just had to include it. We timidly phoned Jo Alexander and explained. This was a native plant. Could we add it to the key and put a photo and a paragraph somewhere? Anywhere? Please? After a pause, Jo said, "You really want to give it two pages, don't you." Well, yes. "I'll find a way. Get the text and photos to me as quickly as you can." And so Carex lacustris made it into the book.

We can honestly say that the second edition of the Field Guide to the Sedges of the Pacific Northwest is up to date. We hope that it will be at least as useful as the first edition for those dealing with the sedges of this area.

STEPHEN SHARNOFF AND LICHENS OF CALIFORNIA

Sharnoff, Stephen. May 2014. A field guide to California lichens. Yale University Press, New Haven (www.yalebooks.com). ISBN 9780300195002 [soft cover] xvi+ 405 p. Price: US$32.50.

Lichens of California (1988) by the late Mason E. Hale, Jr. (1928-90) & Mariette Cole (see Taxon 38: 624) had been the standard work for two and a half decades. Number 54 in the venerable series California Natural History Guides, the book measures just 204 × 124 mm (paperbound) and has only vii, 254 pages and 12 color plates. The text has 40 B&W maps and 71 B&W drawings and photos. Hale & Cole keyed and described some 350 species of lichens, including 48 common species illustrated in color, and 40 widespread species each plotted on a county map of California, one dot per county showing presence. Much of the photography is marginal in quality. Moreover, 25 years is a long time, especially nowadays when the molecular clock keeps ticking away new taxa.

In October 2001 Yale University Press published Lichens of North America by Irwin M. Brodo, Sylvia Duran Sharnoff (1944- 98) & Stephen Sharnoff. Stunningly and lavishly illustrated with 939 color photos (924 numbered, 15 unnumbered) mostly by the Sharnoffs, this work was dedicated to Hale (and three other persons) and has been highly acclaimed (for review see R. Schmid, Taxon 50: 1290). However, the book is too hefty, bulky, and expensive for field use: about 3.5 kg, xxiii, [i], 795 pages, 287 × 252 × 58 mm (hardbound, ISBN 9780300082494), $69.95 then, $135.00 now (oh inflation!). The 636-page taxonomic part has extensive keys to and comprehensive descriptions of 805 species of lichens, with mention of another 700 species in the keys or notes. Most species have Flora-of-North- America-style thumbnail maps.

Here then were the makings for a new field guide to the lichens of California "that would reflect recent name changes, have good [i.e., superb] color illustrations, and include more crustose species" (p. xiii). Sharnoff choose not to provide "keys or extensive descriptions of microscopic characteristics, choosing instead to use the space to include more species" (p. xiv). An appendix lists "recent name changes": 35 specific, 4 generic. There are no range maps.

Shirley Tucker's (2014) checklist claims for California 1869 species and infraspecific taxa compared to 1575 in her & Ryan's (2006) compilation (see entries below under "Floristics …"). I had to search a bit to find the number of lichen species Sharnoff treats. The number, "some 500," appears rather coyly on the front cover flap. He notes that his guide "includes most of the 'macrolichens' (larger lichens, not crusts) that one is likely to encounter in the state, plus a substantial sampling of the more common and distinctive crustose species" (pp. xiii-xiv).

I really do not miss the nomenclatural authorities, which could have been included after the taxa without lengthening the book; authorities can be found in Tucker's 2014 checklist. I do miss, however, the dot-distribution county maps for selected taxa. Such maps would be at the top of my wish list for inclusion in a second edition.

Sharnoff's Field guide to California lichens is superbly and spectacularly done. It certainly will be a big hit! Coda: I have seen used copies (in "acceptable" condition, the lowest quality) of the paperback of Hale & Cole's Lichens of California, originally $14.95 new, selling on Amazon and elsewhere for thousands of dollars. For example, on 21 May 2014 www.addall.com, which searches over 40 sites, including Amazon, Alibris, and Abebooks, and Half.com (see R. Schmid, Taxon 49: 853), showed copies of Hale & Cole selling from $9.19 to $4,873.47 postpaid. I cannot fathom why anyone would pay much more than $25.00 for the book now, but, if so, he or she may also be interested in buying a certain bridge in New York City.

Subscriptions: http://victoria.tc.ca/mailman/listinfo/ben-l.
Send submissions to aceska@telus.net

BEN is archived at http://www.ou.edu/cas/botany-micro/ben/