| BEN |
| BOTANICAL ELECTRONIC NEWS |
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| ISSN 1188-603X |
| No. 487 February 28, 2015 | aceska@telus.net | Victoria, B.C. |
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This issue of BEN is dedicated to
at the occasion of his 80th birthday on March 1, 2015
See also related photographs by David Denning & others at http://bomi.ou.edu/ben/487/ben487_bell.pdf
I first knew Marc when I was still a kid. His mother, Mrs. Bell, was one of the stalwarts in the Victoria Natural History Society, and as a member of the Junior group, with Freeman King, I and some of the other juniors used to follow Mrs. Bell around and watch as she set up her amazing (then) state-of-the-art recording equipment to document bird calls and other sounds of the forest at Francis Park and other locales around Victoria. Freeman King, or "Skipper" as we called him, was a tremendous fan of Marc Bell, and my parents knew Mrs. Bell and Marc too from their association in the VNHS. So, I remember going with my dad to hear a lecture by the great Dr. Vladimir Krajina when I was in grade 11 or 12, and that talk was introduced by Marc. Then, as I was finishing grade 12, somehow a position opened up for a student to work in the UVic Herbarium under Marc's supervision. I didn't realize until much later that my parents, scheming with Freeman King, and I guess Marc too, created that job for me. I was already totally immersed in botany by then, so learning the ins and outs of the herbarium, collecting plants, vascula and plant presses, was just totally fun. At that time the wonderful Mr. T. R. Ashley was there, an amazing quite elderly Welsh botanist, who used to wear a long black apron and always seemed to have a magnifying glass in his hand. I soon became fast friends with Anthea Fisk (soon to be Anthea Bryan; Anthea and her husband Jim have remained dear friends to this day). There were others there in the herbarium: Brian Davies, Alan Keller, Wendy Armstrong, Joan Hett, all wonderful and devoted plant collectors, all mentored by Marc.
I was to work in the herbarium for three successive summers, and in the last one Bob, then my fiancée, now my husband of almost 46 years) was hired on, and the next year he worked for Hans Roemer as a field assistant, still under Marc's overall supervision. Majoring in Botany, through the Biology Department, I had the privilege of taking Plant Taxonomy and Natural Resources Management courses with Marc, and then, when it came to doing my honours thesis research, he consented to be my supervisor. He was a wonderful teacher. Kind and interesting, inspiring. I remember he would challenge us all the time. One of our exam questions in the Plant Taxonomy course was: "The Ideal Plant Classification System: Possibility or Pipedream?" Of course, we had to talk about what a taxonomic system was, the different ways and criteria by which plants were classified, including genetic, chemical and numerical parameters. It really made us think.
When I started my honours research, I had been torn between studying ethnobotany and studying the issues around pollution and environmental quality, which were, of course, topics of great interest to Marc. I started off with the idea of doing some kind of observational project looking at the relationship between lichens and air pollution, since we were suffering from the smoky pollution from the Victoria plywood mill at that time. However, I just could not seem to come up with a good research design for this topic. I remember well going to see Marc to tell him I just couldn't make a "go" of it. Marc looked me in the eyes and said, "Look. What do you really want to study?" I answered, "ethnobotany," and he said, "Well, go for it!" or words to that effect. That was all the encouragement I needed, and I ended up working with WSANEC and Hul'qumi'num elders, especially Christopher Paul of Tsartlip, documenting the names and relationships of people and plants for the Coast Salish Peoples of Vancouver Island. Bob and I were married the week after we both graduated, in 1969, and set off on our honeymoon, ever grateful to Marc for his constant encouragement, yet always challenging us to think carefully about the issues and circumstances surrounding our work. By the time we had returned from our honeymoon, Marc had arranged for my honours thesis to be published in the journal Economic Botany. After we left to go to UBC, with Marc writing a very kind reference letter for me which I showed to Roy Taylor, who became my graduate supervisor, Marc and his family remained friends. He was, and is, just that kind of person, ever encouraging, ever positive. I know that what he did for Bob and me he also did for so many others; he inspired all of us and helped us launch into whatever our botanical or other careers would become.
In the 1970s, Marc was one of the key founders of what is now the School of Environmental Studies, where my position at the University of Victoria has been since 1990. If we think of the multiplier effect of Marc Bell – all those students he inspired, the UVic Herbarium he developed, the program of Environmental Studies that he helped found, and later, the Elderhostel Program that he lead to teach mainly seniors and retired folks about the wonderful world of botany – it is just remarkable. What a positive force he has been in the world, in his quiet and unassuming way, and so many of us have benefited from his gifts.
Marc was a young 32 year old professor at the University of Victoria when he hired me, virtually 'sight-unseen'. Not only that, he brought me and my new wife to western North America from the other side of the globe. How trusting!
In 1966 Marc had spent his first sabbatical visiting individuals and institutions in Europe that represented the classical schools of plant ecology, called "Pflanzensoziologie" in German and plant sociology in English. Not only during his own university education under Dr. Vladimir Krajina, but even more so during his European tour, he became a 'convert' to this approach utilizing a multitude of ecological 'releves' and meticulous tabulation and sorting of these in tables as a basis for vegetation classification. Was I just lucky to be one of the students of Dr. Reinhold Tüxen, then a German authority for these methods? We never met face to face when he was with Tüxen and elsewhere in Europe, but started corresponding and he eventually offered me a one–year assistantship in Victoria.
When we arrived we were most impressed by this young professor, being accustomed to gray-haired, unapproachable authorities from our own university education We soon learned with amazement that we could address him by his first name instead of "Herr Professor".
Marc was a man of action and very generous, helping us in every way, first to smooth our way through immigration bureaucracy oversees, and then with housing and other necessities when new in Victoria.
In my ecological studies Marc gave me a lot of freedom, but was always at hand when I needed practical advice or felt some principles or methods had to be discussed. Marc had a lot of what I would call 'ecological common sense'. As I came reasonably prepared to put names on most of the new trees, shrubs and other forest plants, there was only limited botanical tutoring needed. But I remember well that I was somewhat overwhelmed by the non-vascular plants and that Marc managed to completely put me at ease by pointing out that there are only a limited number of bryophytes growing on the forest floor (most of which he named to me then and there) and that they could be further divided in easily handled groups by distinguishing those that grow on mineral soil as opposed to rotten wood and other organic layers.
After a year of mostly gathering data in the field Marc very generously suggested that if I wanted to work towards a PhD I could incorporate the data thus far collected as his research assistant. He also pointed the way for me to apply for a federal government fellowship, worked closely with me to write the application, and guided it through the channels to its eventual success. Marc was as skillful with words, spoken and written, as he was with ecological principles. This was particularly helpful when it came down to my writing of the dissertation and what I retained from this time has certainly benefited me for the rest of my career.
Thank you Marc, and happy birthday!
In 1966, Marc Bell was traveling through Europe and one of his last stops there was in the Botanical Institute in Pruhonice near Prague where he was a guest of Dr. Emil Hadac. I was a Ph.D. student there working on classification and ecology of wet alder forests (Alnetea glutinosae). One nice afternoon I was walking to catch the bus home. My head was full of the mean (average, not nasty!) similarity coefficients when I ran into Emil Hadac, who offered me a ride home. Emil Hadac lived in Prague walking distance from our apartment. I accepted his offer, but was rather intimidated by the foreigner with him who was strolling among rhododendrons in front of the Pruhonice Castle. It was Marc Bell. Emil drove us home and I flooded Marc with my great ideas about the similarity measures in my broken English, English that he could not understand at all. His mind was also still fully occupied by the accident in France, where thieves had stolen his passport. It was not only the passport, the thieves also stole his tie. Passport yes, but Marc could not understand why they had to steal his tie as well. That question occupied Marc about as much as my coefficients occupied my mind.
I knew the answer right away, but was not brave enough to tell it to Marc when I hitched the ride with him and Dr. Hadac. In North America, the broad ties with the so-called "grandfather's knots" were in fashion in the 1960's, whereas in Europe, it was a narrow tie with a narrow, flute-like knot that was in the fashion there. The thieves had to steal Marc's tie, since his passport without a matching tie was entirely useless to them.
Happy birthday, Marc!
What began as the B.C. Vegetation Working Group in 1986 became Botany BC – an annual meeting of botanists that is still held at various locations across BC. Back in March 1986, Marc Bell and a small group of vegetation specialists met at the Pacific Forestry Centre to discuss the formation of a B.C. Vegetation Working Group. Marc had been involved in earlier years with a group of senior ecologists who discussed classification approaches and initiated the development of a standard BC data collection form. Marc Bell and Del Meidinger felt it would be useful to start up a new coordinating and education group to foster communication among individuals involved in various aspects of the description and management of vegetation. The group that met agreed and it was decided to hold an annual workshop that would bring vegetation specialists together in a field trip/annual meeting and possibly symposium format.
The first workshop was held at the University of Victoria on 22nd April 1986 – the agenda was mostly about the formation of a working group and to keep with the objectives that Marc and other felt important to such a meeting, it was followed by a field trip to some local Garry oak stands and rocky shore communities.
The first large meeting was held at the Cowichan Lake Research Station, Mesachie Lake, in October 1986. The theme of that meeting was classification: vegetation, habitat, estuarine, grasslands, etc. The next meeting was again at the Cowichan Lake Research Station in April 1987. The talks at that meeting dealt with two main topics: 1) rare and endangered plants and ecosystems, and their protection, and 2) vegetation succession and seral classification. Most of the speakers were BC specialists, however, a some speakers were invited from adjacent jurisdictions: Dr. David Murray presented Rare plants and their protection in Alaska; John Gamon talked about The rare plant inventory in the State of Washington; Dr. Leslie Viereck, presented Forest succession research in interior Alaska; Dr. Paul Alaback spoke about Research in plant community dynamics in southeastern Alaska; and Dr. Roger del Moral presented Patterns of vegetation recovery on Mt. St. Helens. The field trip was to the Honeymoon Bay Ecological Reserve. It was also at this meeting that it was decided change the name of the group to Botany B.C., to avoid confusion with the "B.C. Vegetation Management Committee" – a group of silviculturists interested in vegetation control.
After the initiation of this group in the spring of 1986 by Marc Bell and Del Meidinger at the University of Victoria, there have been 29 annual meetings throughout BC. The focus broadened over time from a meeting of government staff and academics to a meeting that included amateur botanists and consultants. As in the early years, experts were often invited from other jurisdictions and from one of these – the attendance of Art Kruckeberg in 1996 (Fort St. James) Botany WA was formed.
[Editorial Note: In spite of the fact that "BOTANY BC" is an acronym for "Botanical Association To Accomplish Nothing Yearly", we accomplished a lot! Thank you, Marc, for your vision!]
Dedicated to Marc Bell on the occasion of his 80-th birthday
Utricularia ochroleuca R.W. Hartman, described in 1857, did not gain recognition as a species distinct from Utricularia intermedia Hayne for some 57 years by European botanists. Since it is a species regarded as a vegetative apomict, purported of hybrid origin with Utricularia minor L. and U. inermedia parentage, persisting and dispersing via turions (P. Taylor 1989), it has often been mentioned in passing as a hybrid that might be encountered—as Barre Hellquist and I did in our Aquatic and Wetland Plants of Northeastern North America (Crow & Hellquist, 2000). On the other hand, when Adolf Ceska and Mark Bell (1973) pursued a study of Utricularia of the Pacific Northwest they gave full attention to a discussion of U. ochroleuca, including description and distribution, along with an excellent illustration prepared by Oluna Ceska, and wrestled with the problem of Asa Gray's U. occidentalis, described from Washington.
In preparing the taxonomic treatment of Utricularia for my manuscript of Lentibulariaceae for Flora North America North of Mexico (Crow 2014) I, too, had to come to grips with the identity of Utricularia ochroleuca. And while the species can be readily distinguished from U. intermedia and U. minor when these are flowering, all too often U. ochroleuca occurs only in the vegetative stage. Therefore I was intrigued when I can across a study of Utricularia in Scandinavia by G. Thor (1988) where he carried out a detailed investigation of the taxonomic value of the quadrifid trichomes that occur within the bladder-traps. Perhaps there is hope in utilizing these quadrifids to resolve identifications of the all-too-frequent sterile specimens of U. ochroleuca.
As a result of his study of Utricularia of the Nordic region, G. Thor (1988) determined that the U. ochroleuca complex actually consisted of two taxa, and described a new species, Utricularia stygia Thor, with both species distinguished from U. intermedia by having a spur which is half as long as the lower corolla lip and oriented at an acute downward angle from the lower lip, in contrast to U. intermedia having a spur appressed to and only slightly shorter than the lower lip. Additionally, while all three have dimorphic vegetative bodies, U. ochroleuca and U. stygia have traps borne on both the flat green ("Wassersprosse") shoots as well as the colorless shoots ("Erdsprosse"), whereas in U. intermedia, the bladder traps are typically borne only on the colorless shoots (very rarely with a few occurring on leaves).
Thor (1988) distinguished Utricularia stygia from his more narrowly defined U. ochroleuca as having slightly larger, dark yellow flowers with a slight reddish tinge, and the lower corolla lip almost flat, measuring 9¬–11 × 12–15 mm; Utricularia ochroleuca in the narrow sense having smaller paler yellow flowers, and the lower lip almost flat, but with the margins later becoming slightly deflexed, measuring ca. 8 × 9 mm. However, as it is not uncommon to encounter these two taxa occurring only in the sterile/vegetative state, identification is further complicated. Thor distinguished Utricularia stygia by having flat leaf segments with 2–7 marginal teeth bearing bristles, whereas U. ochroleuca was characterized as having leaf segments with 0–5 teeth with bristles—the overlap in this character being rather considerable. Furthermore, Thor's detailed examination of the shapes of the quadrifid glandular hairs within the bladders proved useful as a taxonomic character, and he especially regarded the angle between shorter arms as being diagnostic—U. stygia: quadrifids somewhat X-shaped, with small arm angle (30º–)52º–97º(–140º), mean 72 ±22; U. ochroleuca: with upper small arms strongly divergent, forming an angle (117º–)146º–197º(–228º), mean 171.2º ±25.4º. The quadrifids of U. intermedia are quite distinct in being H-shaped (the arms parallel, thus forming almost no angle); the quadrifids of U. minor are also distinctive, with the short arm so greatly divergent that the angle typically exceeds 180? and appear reflexed.
B. J. Plachno and L. Adamec (2007), citing the rarity of flowering in both Utricularia stygia and U. ochroleuca, as well as the unreliability of the number of teeth along the leaf segments as diagnostic, embarked on a study of the differentiation of the two taxa based on the quadrifid glands for plants of the Czech Republic (both taxa having "endangered" status in that country). Of the various parameters studied (including lengths of long/short arms and all possible angle measurements) they concluded that the only statistically reliable criterion distinguishing these two species in the sterile state is the angle between the shorter arms. But they also found that, while the mean value in angles between the shorter arms (74º ±22º) of U. stygia in populations from South Bohemia fell neatly within the range for the taxon as presented by G. Thor (1988), the same was not true for U. ochroleuca. For the latter taxon they found that the angle means (110º–135º) between the short arms of plants were much lower for South Bohemian populations than the mean angle for typical Nordic plants (171º ±25º) as reported by Thor (1988). They also noted that Thor distinguished two variants within his concept of U. ochroleuca, a typical one for the Nordic countries, and a second variant in Europe outside Scandinavia with smaller quadrifid glands and a distinctly smaller angle between short arms than observed for the Nordic plants.
Eric Schlosser (2003), a European, was the first to report the occurrence of Thor's new species, Utricularia stygia, in the United States—Willow Lake, Plumas Co., California. In an effort to understand "what is U. stygia?" he examined quadrifids. He noted that the values of means of angles between short arms of quadrifids in plants from East Germany and from South Bohemia to be "intermediate" between U. stygia and U. ochroleuca with respect to the values given by Thor (1988). Schlosser added that:
"While Thor chose a form of U. ochroleuca s. str. common in northern Europe for his description, the type specimen of U. ochroleuca sensu str. used by Hartman, and the form that may occur more often elsewhere in Europe, are reported to have typically smaller values for the quadrifid angle S [short arms], and more teeth along the margins of the terminal leaf segments. Thus it is less different from U. stygia and the casual observation of a few quadrifids is not foolproof." [My emphasis]
In fact, Plachno and Adamec (2007) suggest that although there seems to be less variability in "good" Utricularia stygia, nevertheless "...there are small patches of quadrifid glands with angles reminding U. stygia," and indicate that, as a consequence of overlaps in the angles of both taxa, it is necessary to examine a minimum of five bladder traps and at least 40–50 quadrifids, focusing especially the maximum angles (U. stygia: mean angle less than 85º, maximum angle less than 115º; U. ochroleuca: mean angle greater than 100º, minimum angle 70º, at least 10% angle greater than 120º or 130º). The occurrences in North America of Utricularia ochroleuca have been rare and widely scattered. In addition to mapping the range of U. stygia in Europe, where numerous populations of both taxa occur, Thor (1988) also cited herbarium specimens from the Gray Herbarium of Harvard University from Nova Scotia (St. Paul Island), and from Alaska, but since his study related to the Nordic countries, he did not otherwise specifically address the occurrence of his new species, Utricularia stygia, in North America.
In reexamining the specimens from the Gray Herbarium studied by Thor, as well as numerous additional specimens of the Utricularia ochroleuca complex, I especially focused on the value of the quadrifid trichomes as a diagnostic character—after all, vegetative plants are more likely to be encountered. The results were mixed. Based on quadrifids, six specimens fell within Thor's concept of U. stygia, with a few additional samples that appeared intermediate between "U. stygia" and U. ochroleuca sensu str. in quadrifid morphology. And yet, there is something "fishy" about Willow Lake, California, wherein Schlosser reported U. stygia. I examined 5 specimens from that lake, and all had quadrifid morphology of "U. stygia," but 2 sheets collected by Barry Rice (DAV) had turions that were very setulose and the spur of the corolla was constricted—both characters of U. intermedia—as well as a general plant habit and leaf morphology strongly suggesting U. intermedia, and I so annotated those specimens. And Rice notes on his specimen label (Rice BR060704 DAV): "Numerous plants growing with U. minor. Plants at this site continue to be taxonomic troubling."
But what about Asa Gray's Utricularia occidentalis, from Falcon Valley, Washington Territory, long treated as a synonym of U. ochroleuca? Of particular importance to me was the status of quadrifids from specimens collected by W. N. Suksdorf from the type locality in Klickitat County, Washington. My examination of Suksdorf 11472 (DS, now housed at the California Academy) resulted in a quadrifid mean of 105º— "intermediate" between U. stygia (72º ±22) and U. ochroleuca sensu str. (171º ±25) per G. Thor's criteria, as well as by the means established by B. J. Plachno and L. Adamec (2007) for U. stygia (62) and U. ochroleuca (126.4). Thus, based on my studies the type specimen fits within U. ochroleuca in the narrow sense, although on the low end of the overall range for U. ochroleuca.
Should Utricularia occidentalis been determined conspecific with U. stygia, then the name Utricularia occidentalis would have had nomenclatural priority—Asa Gray having described the species in 1883 (Proc. Amer. Acad. Arts 19: 95). Interestingly Adolf Ceska (pers. com.) related to me: "I wonder if Utricularia stygia vs. Utricularia ochroleuca are not [merely] two different products of hybridization, one being the male of U. intermedia × female U. minor and the other one just the other way around." I have wondered that myself. And with the Willow Lake locality for "U. stygia" having an abundance of U. minor (per Barry Rice's specimen label), as well as a presence of U. intermedia, the latter just might be true. Resolution will surely require applying molecular tools. [Editorial note: Chromosomal DNA?] Meanwhile, it appears to me that although Utricularia stygia may be worthy of recognition at some taxonomic rank, perhaps at the varietal or possibly the subspecific level, yet at the species level it is at best a "cryptic species." Therefore, until some more definitive study can be conducted, it is more practical to treat this group taxonomically in the broader sense, with U. ochroleuca having nomenclatural priority and U. occidentalis and U. stygia as synonyms.
Treating Utricularia ochroleuca in the broader sense, it is a taxon of bogs, boggy meadows, and marshes, often occurring in shallow water, but especially tending to remain vegetative if in deeper water of streams and lakes; its distribution is circumboreal: Greenland; Alta., B.C., ne. Man., N.S., nw. N.W.T., Ont., nw. Que.; Alaska, n. Calif., c. Colo., Mont., w. Oreg., w. Wash., nw. Wyo.; Eurasia. Although still regarded as rare, it may occur more frequently than presently known, but overlooked because of its penchant for being vegetative. Just last month I confirmed a second population (both non-flowering) discovered in Montana within the last 2 years.
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