ISSN 1188-603X

No. 488 March 5, 2015 Victoria, B.C.
Dr. A. Ceska, 1809 Penshurst, Victoria, BC, Canada V8N 2N6



Botany BC 2015 will take place July 2nd - July 5th 2015 in and around Elkford BC in the beautiful Elk Valley in the southeastern corner of BC! Details will be posted to the Botany BC website at:

Since we changed this year's location from Clearwater to Elkford in October we are a little further behind in the planning process than we had hoped but then again, that's nothing really new. It does mean that there isn't much more detail than what is included here at the moment but a number of negotiations are underway so we expect that the draft program and registration should be up in March.

Please feel free to direct your questions to


From: Dieter Mueller-Dombois, Emeritus Professor of Botany & Ecology, University of Hawaii at Monoa, Honolulu, HI.

Dedicated to Marc Bell on the occasion of his 80th birthday

[Editorial Note: Prof. Dieter Mueller-Dombois was the first one who answered my call for articles dedicated to Marc Bell. Unfortunately, he sent it to my old email address that does not work anymore. Apologies to Dieter, Marc, and all of you who read BEN! Adolf Ceska]

Marc Bell was my field research assistant when I was starting my PhD research in 1955 in Nanaimo River Valley on Vancouver Island. We were housed together in a cottage near the first lake provided by the Comox Logging Company. The Company also supplied us with a pickup truck and food, a wonderful support package received through the Company's CEO Mr. Mulholland, himself an engaged professional forester. Our Prof. Dr. Vladimir J. Krajina had asked for this great arrangement. My wife did the cooking. Marc helped wherever he could, even looking after our new-born baby and first daughter Christine, born 1955, now in her 60th year.

On many sunny mornings, I remember Marc singing his standard song " oh what a beautiful morning, oh what a beautiful day, everything is going my way…" Marc was a passionate fly-fisherman. In our free time, he tought me the skill of fly-fishing. We often drove evenings after work to the Fourth Nanaimo Lake, where we caught plenty of Rainbow trout, a great adventure. Marc, at that time still a forestry student at UBC, proved to be a great field assistant and supportive companion. We did over 100 vegetation samples [relevés] together, dug an equal number of soil pits for soil profile analyses, laid out a good number of square-meter sized seed traps to study seed distribution patterns in the logged and burnt cutover areas of the Douglas-fir forest in Nanaimo River Valley along the four lakes and its neighboring mountain slopes. In the relevé sampling one of us took the notes, the other called out the species names. We collected a good number of plants. A separate task was counting the tree seedlings in subplots of our 400 square meter relevés. We also monitored seedling growth and survival and measured soil moisture and temperatures with max./min. thermometers and evaporation from black & white Livingston atmometers.

While doing the weekly monitoring rounds, one day a small catastrophe happened in the back of our pickup truck, an explosion with fire. The reason was that one of the gallon glass bottles had only a small amount of distilled water left that functioned in focusing sunrays like an enlargement glass on igniting paper. We always kept two glass bottles in a cardboard box padded with newspaper with distilled water for weekly refilling the atmometers. This explosion was one of two small accidents that happened to us. The other was the shear breakoff of the right front wheel of our heavily used and handed down pickup truck, when we were driving up the logging road at Echo Lake.

At that time in the 1950s, our forestry curriculum was still very much devoted to forestry as a business, i.e. use of forest products and how to get the forest out, including slash-burning as clean-up job after logging to prevent larger fires, and not so much with the task of how to grow the forest back. Marc and I had many discussions on that, and I persuaded him in the direction of forest ecology, as a worthwhile and appropriate professional endeavor, particularly in British Columbia. The result of this "friendly persuasion" was for him to follow my footsteps and to go for a PhD in forest ecology under our beloved and inspiring professor Vladimir J. Krajina. Marc then followed through with his decision and acquired his PhD in 1964 with a dissertation entitled "Phytocoenoses in the dry subzone of the Interior Western Hemlock zone in British Columbia" UBC_1964_A1 B4.pdf .

In the following years, after Marc became a botany/plant ecology professor at Victoria University on Vancouver Island, we had limited contact, at three occasions only. He invited me as outside examiner once at the PhD examination of one of his students, Adolf Ceska. I believe it was in 1978. We once more came together in 1993 for celebrating the life of our professor "Kraj" [as Marc called him for short] who had passed away, and a third time in Hawai'i at a professional meeting also in the 1990s, which Marc combined with an ocean fishing trip to Christmas Island.

Both of our pathways became individually so involved that we lost contact thereafter. But my memory of Marc as an early professional friend, with the great time we had together on Vancouver Island, remains alive.


From: Judith Dean Godfrey, Ph.D.

Details from Dr. Wagner's description: Wagner, David H. 2014. Guide to the Liverworts of Oregon. Northwest Botanical Institute, Eugene, Oregon, U.S.A. ( ); E-book, HTML document, 593.8 megabytes on disk, 860+ illustrations, mostly colour photomicrographs. ISBN 978-0-9906193-0-7. Cost: $50 for initial purchase, $10 for upgrade at any time [for future revisions]. Introduction with instructions for navigating using web browser, illustrated dichotomous keys to 170 species, shortcuts to distinctive species, individual species pages, illustrated glossary, literature references, genus index, species index, and classification overview with illustrated exemplars.

What exactly is this Guide? As Wagner says, it is neither a textbook, nor a manual, but an E-book, an HTML document. The main folder, "Oregon Liverworts" is to be loaded onto one's computer hard drive, after which clicking the first file (ahepkey.htm) opens it into the browser. If, at this point, mention of HTML, E-books, and opening in a browser sound a bit intimidating, it should not be. I have to confess here that I had my own technical questions, and Dr. Wagner kindly provided the following information: "This Guide works on any computer, even old computers with outdated software. It works with any browser. If a computer can use a website on the internet, it can use this Guide." If this computerized approach still seems a bit strange after all the times spent "sitting down with a good book" with liverwort keys, the information that comes with the Guide says that printing one hard copy is permitted. A complete hard copy print-out of all the pages as they appear on a computer screen, with small thumbnails of the images, would use approximately 400 sheets of paper.

Once the Guide is loaded onto your computer, however, its General Key, and other features are quite easy to use. Open the main file (ahepkey.htm ). This is the home page and navigation hub. Subsequent pages have links back to the home page, along with back-buttons on each page to make navigation easy.

As for the geographic area covered by the Guide, the focus is on liverworts of western Oregon. Wagner says that coverage of eastern Oregon hepatics is reasonable; coverage beyond the state, to California and north to British Columbia, is less complete. The intended audience of this Guide consists of intermediate to advanced students of bryophytes. It is assumed that the user has some training in general botany.

Main Features:

The main reason for using this Guide is to identify liverworts, so the central feature is the General Key. This is a familiar dichotomous key, with each couplet presenting a choice based on which feature(s) best fit the plant being identified. In this electronic key, there is only one couplet per page. Photos, in particular photos of microscopic features, are used to illustrate each of the two choices in a couplet. The photos are thumbnails, and clicking on each one will enlarge it one or two times; the enlargements are quite useful. Throughout the Guide, it is good to see so many photos of oil-bodies, a very important microscopic feature for identification of species. Oil-bodies can be observed in leaf cells in freshly-collected liverworts, but are often lost as the plants dry. Dr. Rudolf M. Schuster, preeminent hepaticologist and author of The Hepaticae and Anthocerotae of Eastern North America: East of the Hundredth Meridian and a great many other works on Hepaticae, always stressed the importance of studying fresh specimens with intact oil-bodies.

Once a species is reached as an end-point of keying, the helpful species information provided includes some synonyms, special conservation status, recognition hints (morphological features), distribution, comments, and additional illustrations (photos or drawings). I would prefer to see the notes on species habitat and associated species separated out from notes on distribution in the Guide area. Photos for some species descriptions could use additional labels to specify which specific feature one's attention is being drawn to, e.g., for Calypogeia azurea, labels could include "oil-bodies", "underleaves", and "gemmae", as is done elsewhere for other species.

The Guide includes a very good Glossary, clear and well-illustrated with photos (again, thumbnails) and some line drawings. The Glossary is good for quickly checking a term or looking at a photo illustration while keying out a liverwort. The description of "succubous" as "like a Venetian blind turned down" made me smile; a good way to convey that. The Guide includes a Genus Index and Species Index, and all genera and species have clickable links so that a person can quickly find those particular keys. Each genus name links to the first couplet in a key to species of that genus, or where there is only one species in the Guide area, directly to the species page. Links make it easy to compare different species. There doesn't appear to be a general "Search" function. Once one is more familiar with liverworts, the Shortcuts to Distinctive Groups can be used to quickly jump to a particular set of morphological characteristics.

After trying out the General Key a few times, and scanning the Genus Index and Species Index, I looked to see what classification system Wagner uses, and found that it is familiar and follows B. Crandall-Stotler, R. Stotler, and D. Long (2008, Bryophyte Biology, 2nd Edition, eds. B. Gofffinet and J. Shaw, Cambridge). On the species pages and in the indices, the Guide provides some recent synonyms for species names, including some from molecular studies, for anyone interested.

Does the Guide and its General Key work? The first species I test-keyed was a simple one, Marchantia polymorpha. I tried to trick the system with lack of gemmae cups, but the key still led to the correct answer. I subsequently ran through several thallose and leafy species and reached correct end-points for species identification. With the ease of navigation and ability to quickly click links to compare photos and features for different species, it is easy to play with the key when working through it, learning as you go.

While the Guide's General Key works quite well, and is very easy to navigate, there are some points I will mention, not as criticism, but as part of the ongoing conversation toward future revisions and updates.

It is probably true that no two people would construct a key in the same way, and there are some places in the key where I might use different criteria, or a different sequence of criteria. Couplet 3a/3b requires a decision as to whether the leaves are divided less than, or more than, one-half. I found this required decision a bit difficult or confusing when test-keying a complicate-bilobed plant, e.g., a Scapania. The 3a/3b couplet could include an illustration of "complicate-bilobed". Alternatively, "complicate-bilobed" could be used as a main split from "simply-lobed" (or some such wording) prior to 3a/3b, to avoid a jump in couplet criteria from leaf lobes to oil-bodies and back to leaf lobes. Of course, with a bit of experience with liverwort morphology and in using the keys, a person could simply use the Shortcuts to jump to groups of liverworts with complicate-bilobed leaves. In addition, I would suggest starting the text for 3b with "leaves entire", to match the accompanying line drawing of Gyrothyra underwoodiana.

There are likely a few mistakes or glitches in a work of this complexity. For example, there is an apparent mistake on the Schofieldia monticola species page, in labeling of other liverworts mixed in with Schofieldia in the photo. The "L" should refer to the subalpine Lophozia opacifolia rather than L. incisa. The two Lophozia species are correctly described and differentiated on their respective species pages. In the captions for the three photos of single leaves, the references to the letters "S", "L" and "C" have no relevance.

There was one real surprise for me in scrolling through the Guide's Species Index, i.e., the rare species Haplomitrium hookeri from Sutton Beach, Oregon. I wondered, what? Haplomitrium hookeri at sea level, with Phaeoceros and Blasia? In my work on hepatics in British Columbia and northern Washington State, all collections of this species were from high elevations (subalpine and alpine), occurring with such other subalpine liverworts as Schofieldia monticola, Pleurocladula albescens, and Lophozia opacifolia. The loose growth habit in the plant photographed in the Guide differs from my own collections and those made with Dr. Wilf Schofield in our area. In my 1977 Ph.D. thesis (Hepaticae and Anthocerotae of Southwestern British Columbia, I described the plant's growth habit as "fleshy, resembling minute 'Brussels sprouts,' arising from a colorless, rhizomatous basal region". Schofield similarly describes these plants as somewhat resembling miniature Brussels sprouts in his 2002 book, Field Guide to Liverwort Genera of Pacific North America. His illustrations show the growth habit with closely overlapping leaves that led us to this description. I have read that growth habit of this species can vary with conditions, and also that in other locations (e.g., northern Germany, Poland), it apparently may occur at lower elevations. Wagner mentions that the Sutton Beach population has disappeared. Hopefully, someday, more populations will be located to enable additional study of this rare plant's habitat and distribution in the Pacific Northwest.

Overall, it was interesting, fun, and a pleasure to work through this Guide and get to understand it better by trying it out for this review. The Guide is a good way to identify liverworts, especially when used along with the recommended book references. It is an excellent learning tool, for self-study to learn liverwort features, as well as learning to identify species.

Although this Guide focuses on liverworts of western Oregon, it also will be helpful to botanists in other regions who wish to learn about liverworts. A person would just have to work through it to the generic level, learn from the photos, glossary, and other information, and then supplement with information about regional and local species. Liverworts are truly interesting and very beautiful plants with a great diversity of forms, as the photos show. David Wagner has developed this E-book Guide over many years, beginning with typed drafts of the keys, then switching to HTML by about 2000. In this form, the Guide can be easily distributed, and updated, at a cost far less than that of a formally published book. Thus this information will be accessible to a greater number of teachers and their students, and to people who just want to learn about liverworts on their own. This Guide is a welcome complement to the traditional manuals and keys for identification and study of liverworts.

I am grateful to Dr. Geoff Godfrey for helpful suggestions regarding this review

Send submissions to

BEN is archived at