ISSN 1188-603X

No. 300 December 26, 2002 Victoria, B.C.
Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2


This is the last issue of BEN for the palindromic year 2002. For me it was an interesting year. At the end of March I retired from the British Columbia government and returned to the "geobotanical" consulting that my wife and I did more than twenty years ago. In July, I celebrated the first anniversary of my heart attack: the cardiologist told me then that I had a genetic defect and was supposed to die - suddenly and without any obvious cause - when I was about seven years old. Luckily, I lived long enough to see the progress of medical technology. When my heart troubles showed up, the doctors were able to find that extra node in my heart. They "mapped it and zapped it" and I feel better than ever before. This past year I spent more days in the field than in the past five years together. "Study nature, not books," is an aphorism attributed to Frederick Edward Clements, the father of North American plant ecology. I have definitely done my share of studying this year.

This is also the last issue of BEN in the third century of BEN. [Cen-tu-ry: a group, sequence, or series of 100 like things; see also Bob Ogilvie's note in BEN # 100.] This event happens approximately every four years, but only with this issue, the beginning of a new year came together with the beginning of the new century for BEN. At this opportunity I would like to thank all who contributed to BEN with their articles and notes. My thanks go to the Victoria Freenet Association - where the BEN mailing list resides - and to Prof. Scott Russell, who is working hard as a reliable web master in the BEN archives at Oklahoma University. Several editors turned my writing into (I hope) readable language, and occasional reviewers gave me their editorial help with those contributions where I was unable to decide about the contributions' merit or accuracy. Many thanks to you all! I would like to remind all of you who read BEN that I need your articles, notes, announcements, etc. in order to keep BEN going. If you are making any resolutions for the coming year, please, add this one:

This year I will write something for BEN.

Have a happy and most successful new year 2003, and best of luck in keeping your resolutions!

Adolf Ceska


Sad news to all veterans of Botany BC: Al van Ryswyk died in Kamloops on November 19, 2002. Best known to many as "van", Al was born in Castor, Alberta on May 10, 1928. He grew up in Castor, where he rode horseback 5 miles to school every day. In 1945 his family moved to Ladner, BC, and Al entered the University of British Columbia. He graduated in 1949 with a B.Sc. in Agriculture. Upon graduation he took a summer job with the BC Soil Survey (BCSS) in Kelowna, BC. Fortunately by fall, he took an appointment to do soil survey work with the UBC Doukhobor study group that led to his MSA in Soils (1955). In the spring of 1958 he received a permanent appointment to Agriculture Canada, Kamloops (then the Experimental Farms Service). Educational leave to Washington State University culminated in the award of a PhD (1969 - Soils with Henry Smith and Plant Ecology with Rexford F. Daubenmire). van Ryswyk's research contributed greatly to our understanding of the classification, fertility, and productivity of grassland, forest, wetland and alpine soils. Al retired in June 1994 after 37 years of service with Agriculture Canada.

Al was a member of the BC Institute of Agrologists and recipient of a Recognition Award in 1998 for his outstanding leadership and professionalism within the agricultural industry. He was a life-member of the Society for Range Management and president of the B.C. Chapter in 1983-84. In September 2002, he was honored with the prestigious Trail Boss Award by the Society for Range Management for his life-time contribution to range management.

He is survived by his wife Alwilda Audrey (nee Minette) and their two sons, Leonard Albert and John Albert van Ryswyk.


From: David C. Tank, Department of Botany, University of Washington Seattle, WA 98195 []

Recently, the Scrophulariaceae (Figwort family) have been the subject of considerable attention in molecular systematic studies. These studies suggest that Scrophulariaceae as traditionally circumscribed, including approximately 220 genera and more than 3000 species, represent a non-monophyletic assemblage of plants identified by shared characters that may be plesiomorphic or commonly recurring throughout the Lamiales (Olmstead & Reeves 1995; Olmstead et al. 2001). In addition, gene phylogenies from analyses of multiple chloroplast genes have identified a well-supported clade containing the obligately parasitic Orobanchaceae and all of the hemiparasitic scrophs traditionally placed in tribes Buchnereae and Rhinantheae of subfamily Rhinanthoideae (dePamphilis et al. 1997; Nickrent et al. 1998; Young et al. 1999).

The subtribe Castillejinae of tribe Rhinantheae is comprised of hemiparasitic species in six genera distributed chiefly in western North America. Bentham (1846) first proposed the subtribe to include three genera, Castilleja (the largest with species as currently circumscribed), Cordylanthus, and Orthocarpus. Later, three Mexican monotypic genera, Clevelandia, Gentrya, and Ophiocephalus, were also placed in the subtribe based on morphological resemblance to the other three genera of Castillejinae (Breedlove and Heckard 1970; Chuang and Heckard 1986). However, throughout the history of the group, there has been significant difficulty defining clear-cut generic boundaries, especially between Castilleja and Orthocarpus (Eastwood 1909; Keck 1927). Extensive morphological study, including floral morphology, seed and seed-coat morphology, and cytological analyses, lead to a major generic realignment of Castillejinae (Chuang and Heckard 1991, 1992). In a previous revision by Keck (1927), Orthocarpus was comprised of 25 species divided into two subgenera (Orthocarpus and Triphysaria), with the type subgenus (Orthocarpus) further partitioned into three sections (Monosaccus, Castillejoides, and Cordylanthoides). In their reorganization, Chuang and Heckard redefined Orthocarpus to include only nine species corresponding to Keck's section Monosaccus (Chuang and Heckard 1992). Subgenus Triphysaria, distinguished both morphologically (the reduction to one anther sac) and cytogenetically (anueploidy to n=11 from n=12), was elevated to generic status to include five species distributed mostly in California. The two sections Castillejoides and Cordylanthoides were determined to resemble the Castilleja pilosa group in floral structure, pollination biology, seed type, and basic chromosome complement, and thus, were assigned to section Oncorhynchus in the genus Castilleja (Chuang and Heckard 1991). Finally, the monotypic genus Gentrya, distinguished by its uniquely curved corolla, was determined not to differ significantly enough from the expanded Castilleja to warrant generic status, and was reassigned to subgeneric status within Castilleja.

My research has been focused on elucidating the phylogeny of subtribe Castillejinae through the use of molecular phylogenetic methods. In their realignment of the six genera comprising the subtribe, Chuang and Heckard (1991) have proposed an 'intuitive' phylogeny of Castillejinae. However, because of the difficulties present in defining diagnosable characters for many of the genera, the question of whether this new organization represents natural, monophyletic groups remains.

The results of separate and combined phylogenetic analyses from multiple chloroplast and nuclear ribosomal DNA regions suggest that: 1) subtribe Castillejinae is monophyletic, 2) Orthocarpus, sensu Chuang and Heckard, is monophyletic, 3) Cordylanthus is bi-phyletic, however its two subgenera Cordylanthus and Hemistegia are each monophyletic, 4) the largest genus Castilleja is paraphyletic, with the monophyletic genus Triphysaria and the two monotypic genera Clevelandia and Ophiocephalus arising from within it, and 5) Castilleja racemosa (previously Gentrya racemosa) is nested within Castilleja.

From these results, it is clear that the Chuang and Heckard's earlier taxonomic work was quite close to defining natural, monophyletic groups in the subtribe. The revised Orthocarpus, including only nine species, forms a well-supported clade. The two monophyletic subgenera of Cordylanthus, Cordylanthus and Hemistegia, are identifiable based on leaf morphology, but do not together form a clade. Species belonging to subgenus Cordylanthus have leaves with varying degrees of division, while those in subgenus Hemistegia have small, entire leaves. Unfortunately, the monotypic subgenus Dicranostegia, comprised of Cordylanthus orcuttianus, has not been sampled yet. It will be interesting to determine the placement of this enigmatic species with repect to the other two Cordylanthus clades. The remaining genera comprising the subtribe are all derived from within Castilleja, and are allied with the annual Castilleja species of section Oncorhynchus. The four genera Castilleja, Clevelandia, Ophiocephalus, and Triphysaria form a very well-supported clade tenatively referred to as Castilleja s.l. Castilleja s.l., in which the majority of species are perennial, has evolved from a grade of annual taxa represented by the annual members of Castilleja s.l. (Castilleja sect. Oncorhynchus, Clevelandia, Ophiocephalus, and Triphysaria) Cordylanthus, and Orthocarpus. This stands in stark contrast to the conventional wisdom that annuals are derived from perennial ancestors. In their intuitive phylogenetic hypothesis of Castillejinae, Chuang and Heckard (1991) propose up to six independent origins of annual habit, rather than one origin of the perennial habit as these results suggest. These analyses also suggest that the perennial Castilleja clade has undergone a recent and rapid radiation in western North America, as compared to its annual progenitors.


Bentham, G. 1846.
Scrophulariaceae. In A. DeCandolle [ed.] Prodromus 10: 180-586.
Breedlove, D. E., and L. R. Heckard. 1970.
Gentrya, a new genus of Scrophulariaceae from Mexico. Brittonia 22: 20-24.
dePamphilis, C. W., N. D. Young, and A. D. Wolfe. 1997.
Evolution of plastid gene rps2 in a lineage of hemiparasitic and holoparasitic plants: Many losses of photosynthesis and complex patterns of rate variation. Proceedings of the National Academy of Sciences, USA 94: 7367-7372.
Chuang, T. I., and L. R. Heckard. 1986.
Systematics and evolution of Cordylanthus (Scrophulariaceae-Pediculareae). Systematic Botany Monographs 10: 1-105.
Chuang, T. I., and L. R. Heckard. 1991.
Generic realignment and synopsis of subtribe Castillejinae (Scrophulariaceae-tribe Pediculareae). Systematic Botany 16: 644-666.
Chuang, T. I. and L. R. Heckard. 1992.
A taxonomic revision of Orthocarpus (Scrophulariaceae-tribe Pediculareae). Systematic Botany 17: 560-582.
Eastwood, A. 1909.
Synopsis of the Mexican and South American species of Castilleja. Proceedings of the American Academy of Arts and Science 44: 563-591.
Keck, D. D. 1927.
A revision of the genus Orthocarpus. Proceedings of the California Academy of Sciences Series 4 16: 517-571.
Nickrent, D. L., R. J. Duff, A. E. Colwell, A. D. Wolfe, N. D. Young, K. E. Steiner, and C. W. dePamphilis. 1998.
Molecular phylogenetic and evolutionary studies of parasitic plants. In P. S. Soltis, D. E. Soltis, and J. J. Doyle [eds.] Molecular Systematics of Plants II, DNA Sequencing 211-241. Kluwer, Boston, Massachusetts, USA.
Olmstead, R. G., and P. A. Reeves. 1995.
Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndhF sequences. Annals of the Missouri Botanical Garden 82: 176-193.
Olmstead, R. G., C. W. dePamphilis, A. D. Wolfe, N. D. Young, W. J. Elisens, and P. A. Reeves. 2001.
Disintegration of the Scrophulariaceae. American Journal of Botany 88: 348-361.
Young, N. D., K. E. Steiner, and C. W. dePamphilis. 1999.
The evolution of parasitism in the Scrophulariaceae/Orobanchaceae: Plastid gene sequences refute an evolutionary transition series. Annals of the Missouri Botanical Garden 86: 876-893.


From: Adolf Ceska []
Dierssen, Klaus & Barbara Dierssen. 2001.
Moore. (Ekosysteme Mitteleuropas aus geobotanischer Sicht.) [Mires. (Ecosystems of Central Europe from the geobotanical viewpoint.)] Verlag Eugen Ulmer, Stuttgart (Hohenheim). 230 p. ISBN 3-8001-3245-1 Price: 59.90 EUR

Ordering information:
Verlag Eugen Ulmer GmbH & Co., Wollgrasweg 41, 70599 Stuttgart (Hohenheim) Germany - e-mail: ; web:

In 14 chapters (the 14th is a list of literature references) the authors cover all aspects of mires. The first chapter ("Mires as an environment/habitat") defines basic terms and processes and gives the key to the chapters that follow. The following chapters deal with hydrology (Ch. 2), palaeoecology of peat deposits (Ch. 3), classification of mires (Ch. 4), peat (Ch. 5), carbon, nitrogen and nutrients dynamics (Ch. 6), main habitat gradients (Ch. 7), vegetation of mires (Ch. 8), overview of vegetation complexes (Ch. 9), mires as ecosystems (Ch. 10), use, threats and protection (Ch. 11), strategies and methods for the protection and restoration of mires (Ch. 12), long-term research and monitoring, and checking of results (Ch. 13), plus literature (Ch. 14).

Klaus Dierssen is a university professor and director of the Ecology Center at the University of Kiel in northern Germany. Barbara, his wife, specializes in botanical and geobotanical photography, ethnology and ethnobotany. I have not read one publication by Klaus (or Klaus & Barbara) that I have not liked. The list includes Most important mire plant communities of northern Europe (in German, published in 1982), Introduction to phytosociology (vegetation science) (in German, published in 1990), Vegetation of Europe (with Barbara Dierssen, in German, published in 1996 - see BEN # 176), and Distribution, ecological amplitude and phytosociological characterization of European bryophytes (in English, see BEN # 273). Barbara and Klaus also published the Botanical travel guide through northern Schleswig-Holstein (in German, published in 1994) that describes about 110 field trips in the vicinity of Kiel.

What can I say about their recent book? Moore fits well into the Diersseniana collection I listed above. The book defines the basic terms, defines the processes, and describes the wetland vegetation of Central Europe in detail. The chapters on protection, restoration, and long-term monitoring discuss conservation and restoration problems. Dierssens' book represents the holistic approach to wetlands, but at the same time it treats the wetlands with great attention to detail. The book resembles Peatlands by Moore & Bellamy (1973) and even has about the same number of pages. It covers the same basic principles, but it goes much deeper into the description of actual vegetation of (Central European) mires. Phytosociology, which was mentioned in Moore & Bellamy in four sentences, is dealt with in Dierssen & Dierssen in the largest chapter (Chap. 8), with almost fifty pages. Several important concepts have developed in "peat science" since Moore & Bellamy (1973) and those new concepts are covered in the recent Dierssens' book (e.g., acrotelm & catotelm). The book is richly illustrated with numerous colour photographs, plentiful graphs, maps and diagrams.

With my poor German I found this book more difficult to read and understand than, for instance, the Vegetation Nordeuropas. I would like to see Moore translated into English, and I wonder if any bilingual readers of BEN would be willing to volunteer to do this. (Please contact me if you are interested in helping.) The potential English edition would open this book to a large English-speaking readership. Although the major portion of the book is devoted to Central European vegetation (Ch. 8 with 49 pp.), many of the plant communities described there are either circumpolar or have homologous community types all over the northern hemisphere. In addition, the prospective English edition should contain a short conspectus of the world floristic classification of wetlands, if something like that can be achieved at this moment.

Summary: In spite of its Central European theme (or maybe just because of it), Dierssens' Moore is an excellent addition to the world literature on wetlands. Job well done, Klaus, Barbara, and the publishers!


CONSTANCEA - University of California Electronic Publications in Botany
Editor: Richard Moe []

Constancea is an online journal that will specialize in presenting material that would benefit from an electronic medium. It supersedes University of California Publications in Botany, which published its first issue June 7, 1902 (A botanical survey of San Jacinto Mountain, by Harvey Monroe Hall, a Master's student of W.L. Jepson), and its last issue, volume 82, in 2001 (Plant diversity of an Andean cloud forest: checklist of the vascular flora of Maquipucuna, Ecuador, by Grady L. Webster and Robert M. Rhode). We anticipate publishing works that are image-rich, that cite numerous specimens and geographic records, that are ongoing projects, that are by their nature hypertextual, or that are particularly timely. Each contribution will be numbered (beginning with 83-to emphasize the continuity with UC Publications in Botany) and announced as it is placed on the Web. It will be marked as to date and version number, and permanently archived electronically as well as on acid-free paper in the UC/JEPS archives. Successive versions, if there are more than one, will also be archived.

We expect that most contributions will come from associates of the University of California, but submissions from outside are welcome as well. Articles in the journal will be rigorously peer-reviewed by at least two anonymous referees.

Current issue: Constancea vol. 83 - Dec. 13 2002 Paul C. Silva 80th Birthday Festschrift [20 articles on algae & bibliography of Paul C. Silva]

Upcoming issues:

VULPIA - Contributions from the North Carolina State University Herbarium [NCSC]
Editor: Alexander Krings [ or]

NCSC is pleased to announce the release of a new online botanical publication Vulpia, Contributions from the NC State University Herbarium, named in honor of the late Dr. William Basil Fox (1915-1952), first curator of NCSC [fox: latin - vulpes].

Not wishing to duplicate existing journal coverage, Vulpia seeks to provide useful scientific information often not published in existing journals for lack of resources or space, such as botanical keys to native and cultivated plants, detailed nomenclatural notes, discussions of synonymy and typification, floristic notes, and lengthy exsiccatae. We hope to serve a broad audience, including professional botanists and amateur plant enthusiasts alike. Manuscripts will be reviewed by the editorial board. Contributors need not be associated with North Carolina State University.

Subscriptions are free for the time being and will be managed through a list-serve. Persons subscribing to the publication will receive notification of new papers as they are released and a complete table of contents of the year's volume at the end of the calendar year.

Volume 1:


Exsiccatae and lists



From: New York Times, December 15, 2002 [abbreviated from]

In the name of science, the United States Forest Service has proposed the experimental logging of half a million acres in two forests in the Sierra Nevada to see how it will affect the habitat of the California spotted owl and the ferocity of forest fires. But skeptical environmentalists are saying the real purpose is simply to give timber companies a chance to cut more big trees on some of the nation's 190 million acres of public land.

The study is to be conducted in the Plumas and Lassen National Forests, two of the 11 national forests that run along the mountainous spine of California.

After a year in which forest fires raged through the West, affecting seven million acres, the administration has been pushing plans to thin the trees in places where years of mismanagement - including the practice of putting out every single fire - have left dense thickets of undergrowth. Often these projects are aimed at protecting small communities at the forest edge. But rarely do they involve cutting so many trees, or such big ones, especially in sensitive wildlife habitat deeper in the woods.

Some trees to be cut are much larger than current forest regulations would allow: in some cases, up to 34 inches [about 85 cm] in diameter.

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