ISSN 1188-603X

No. 458 August 30, 2012 Victoria, B.C.
Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2

DONALD M. BRITTON (1923 - 2012)

From: Dan Brunton

Photo Plate:

On 28 July 2012 a memorial service was held in Guelph, Ontario in honour of Don Britton who died 18 May 2012. Don was widely recognized as Canada's foremost pteridologist, a North American pioneer in the use of cytology in fern classification. He was an acclaimed specialist in the study of numerous Canadian genera including Dryopteris, Polypodium, Gymnocarpium and Isoetes. In her tribute at the Memorial, ex-Britton student and Duke University Botany professor Kathleen Pryer (wearing an appropriately fern themed skirt) described his inspirational leadership as a scientific mentor. She pointed out how "DMB" as she called him was amazingly positive in his encouragement of students to strive for his perfectionist approach to lab procedures but also to never forget the critical importance of understanding the whole organism as a living entity. Blending the test tube and the field, as it were. Daniel Brunton spoke of his experience during the same period, first as one of dozens if not hundreds of field botanists receiving Don's invaluable assistance in specimen identification and later during their in-depth investigations into the genus Isoetes. Both reflected upon Don's exceptional generosity with his time, patience and material support for any and all who shared his curiosity and scientific passions.

As he turned 80 a series of tributes were published to acknowledge his exceptional contribution to Canadian botany (BEN #304 and #305 in 2003) . He was most appreciative of the honour but a bit surprised and overwhelmed by all the fuss. That was so Don. Throughout his long association with the University of Guelph his quiet, unassuming nature was as famous as his quirky and at times cryptic letters. All of his correspondence was composed on an ancient typewritter (he never made the transition to email) with a diagnostic out of alignment letter 'e'. Many of the letters included afterthoughts and addenda scribbled into the margins, usually in red ink. His correspondence was massive, his correspondents legion. No other botanist has ever been so centrally connected to the students of ferns in Canada.

Perhaps most widely known as the co-author with William J. Cody of the 1989 Ferns and Fern Allies of Canada, Don was a prolific author of research papers, usually in collaboration with one or more associates. He produced numerous publications on the cytology, distribution and taxonomy of pteridophytes in North America as well as taxa in Asia and Europe. This production extended into a busy retirement in which he published over 30 papers on Isoetes alone, leading the way in the definition and description of 15 new taxa.

Don received considerable recognition from the scientific community for his many professional contributions. Although too modest to express an opinion on the matter, the tribute he probably was most pleased by was the naming of three pteridophytes in his honour by research associates from Canada, the United States and Europe: Gymnocarpium x brittonianum (Sarvela) Pryer & Haufler, Isoetes x brittonii Brunton & Taylor and Dryopteris filix-mas subsp. brittonii Fraser-Jenkins & Widen. And at the end of the day, isn't it entirely appropriate that there be living memorials to such an exceptional field-based scientist?

[See also BEN # 304 and # 305 dedicated to Don Britton:]


A great opportunity to study mosses and other bryophytes of the Pacific Northwest is coming September 19-21st at the University of Washington in Seattle. The Washington Native Plant Society and UW Herbarium are offering a 3-day bryophyte identification workshop taught by renowned bryologist Dr. David Wagner from Oregon. This workshop is designed to provide amateur and professional botanists with the knowledge and skills needed to develop competence with bryophyte identification. More information about the workshop can be found here:


From: Marcel Rejmánek, University of California, Davis review to be published in Plant Science Bulletin Botanical Society of America

The Jepson Manual - Vascular Plants of California Bruce G. Baldwin, Douglas H. Goldman, David J. Keil, Robert Patterson, Thomas J. Rosatti, & Dieter H. Wilken (eds.) 2012. Second Edition. University of California Press, Berkeley, CA. 1568 p. ISBN 978-0-520-25312-4 (Hard cover) Price: US$ 125.00

The first Manual of flowering plants and ferns of California was published by Willis Linn Jepson, professor at University of California, Berkeley, in 1925. This was an extremely important, but also somewhat overdue accomplishment because seven editions of Asa Gray's Manual of Botany (covering the Central and Northeastern United States) and two editions of Britton and Brown's Illustrated Flora of the Northern United States had been already published at that time (Moore et al. 2010). After 1925, two editions of A California Flora by Philip A. Munz were published in 1959 and 1968. Later, in 1993, the Jepson Herbarium at UC Berkeley, under the leadership of James C. Hickman, produced The Jepson Manual Higher Plants of California (TJM1), effectively starting a tradition similar to Gray's Manual of Botany. This year we got the second edition of The Jepson Manual (TJM2), resulting from revisions and new treatments accomplished by 332 authors and/or editors, produced again by the Jepson Herbarium, but this time under the leadership of Bruce Baldwin.

First, it is interesting to compare numbers of families, genera, and species (total, native, endemic, and alien) included in the three Jepson Manuals (1925, 1993, 2012) and in the summary based on Munz's Floras (Raven 1977):

  Years  1925    1977    1993    2012  
  Native species3727505748444976
  Endemic species    1416152511691315
  Alien species2926541023991

Obviously, the total numbers of families, genera, and species have been increasing almost linearly over the last 90 years. This is partly because of discoveries of native species new to California or even to science, partly because of naturalization of non-native species, and to some extent also because of changes in taxonomy, namely splitting of families and genera into monophyletic units (see below). However, the number of endemic species has been jumping up and down substantially depending on taxonomic delineation of species and improving knowledge of their distribution.

To define the exact number of naturalized (permanently established) species is always difficult (Rejmánek 2007). Authors of TJM2 tried to be more conservative and wanted to include only alien species conclusively naturalized in California. That explains a small drop in the number of alien species compared with TJM1. I can suggest only a few possible corrections: a few species called "waifs" (this is a rather endemic term; "casual" is the term more commonly used Pyšek et al. 2004) are included (Agrostema githago, Cucurbita pepo, Emex australis, Geranium solanderi, Lasiospermum bipinnatum, Triticum vulgare), while some naturalized species (Cuscuta japonica, Danthonia decumbens, Fraxinus uhdei, Geranium yeoi, Melianthus major, Pinus pinea [Santa Cruz Island], Rhamnus alaternus, Rhus lancea, Rytidosperma caespitosum, Solanum mauritianum, Veronica hederifolia) are not mentioned at all or just in keys or en passant in the text. Some species classified as native (Latin names in bold italic) are most likely alien (Galium tricornutum, Landoltia punctata, Phalaris arundinacea, Typha angustifolia). Still, the resulting numbers may indicate recent reduction of the rate of alien plants naturalization in California - a trend anticipated by Rejmánek & Randall (1994, Fig. 1).

In TJM2, the chapter on geologic, climatic, and vegetation history of California was completely rewritten by Constance Millar. The geographic subdivision section was revised and more accurate color maps were provided. There are 272 full plates (compared with 242 in the 1993 edition) illustrating about 80% of native and naturalized species. Genera are somewhat unevenly illustrated (8 of 8 Polystichum species, 18 of 18 Pinus, but only 3 of 12 of Botrychium, 3 of 13 of Eryngium, and 11 of 38 of Lomatium). However, in large and difficult genera (Astragalus, Carex, Eriogonum, Juncus, Lupinus, Mimulus [now in Phrymaceae], Salix), almost all species are illustrated!

As with the TJM1, the index is not all-inclusive, since the book is arranged alphabetically by family and species within each of the eight major clades. In addition, in order to make the volume as short as possible, only synonyms in use since TJM1 are included and can be found in the index and the text; thus species that were moved to different genera in TJM2 can be found by looking in the index. However, if you have only TJM2, you will not be able to figure out what happened to the one of the first species collected in California - Zauschneria californica C. Presl., which was moved to the genus Epilobium in TJM1. In this situation, you have to go to the online Index to California Plant Names; the website URL is provided inside the front cover. In spite of all the editors' attempts at space-saving, TJM2 is 168 pages longer than TJM1. Well, still portable.

Many taxonomic changes were made in TJM2. The authors followed mostly the Angiosperm Phylogeny Group (A.P.G. II, 2003) classification of vascular plant families, which emphasizes monophyletic taxonomy. Goodbye Aceraceae, Asclepiadaceae, Hydrophyllaceae, Lemnaceae; nevertheless, Chenopodiaceae are still kept separate from Amaranthaceae. The relationships of all families, as currently understood, are shown on the back endpaper, along with the page numbers where they are treated. For convenience, within each of the eight major clades, families are presented alphabetically. Genera within families are also organized in alphabetic order. It seems that the prevailing tendency was to split many genera and even some families (for example, the Liliaceae of TJM1 has been split into 12 families, Caprifoliaceae into three, Aster has been broken up into seven genera, Camissonia into nine, Cupressus into three, Gilia into five, Gnaphalium into four, Hemizonia into three, Madia into six, Mitella into three, Polygonum into five, Potentilla into four). This, however, does not seem to be true for the grasses where rather drastic lumping was the rule. Some of these mergers may be justified (Lolium and Vulpia are merged into Festuca; Inda et al. 2008), but others are clearly erroneous (Piptatherum miliaceum is treated as Stipa miliacea!). The classification of some groups of grasses is also not in agreement with recently published volumes of the Flora North America. Moreover, it is not even in agreement with most recent molecular studies of phylogeny in these groups (Romanchenko et al 2010, Hamasha et al. 2012). Yes, our understanding of phylogeny in some groups of grasses is still not complete. However, putting many obviously different species into giant genera was premature. All local floras, databases, and herbaria in California will now follow this provisional taxonomy. In the future, all of that will have to be changed again. A more conservative approach would be less problematic.

Many resources currently available in California made the completion of this manual easier. Just look at The Jepson Flora Project ( or at The Consortium of California Herbaria ( websites to gain an appreciation of this fact. However, we have to realize that this was a difficult time for the taxonomic reorganization of a manual covering so many species. Many major changes in phylogentically based taxonomy at the family level have been accepted since 1993, but the relationships between many genera are still not completely resolved. Obviously, the authors tried their best, and the result is another milestone in Californian botanical literature. Editors and authors should be congratulated on this demanding publication.

Literature Cited

Hamasha, H.R., von Hagen K.B. & M. Röser 2012.
Stipa (Poaceae) and allies in the Old World: molecular phylogenetic realigns genus circumscription and gives evidence on the origin of American and Australian lineages. Plant Syst. Evol. 298: 351-367.
Inda, L.A. et al. 2008.
Dated historical biogeography of the temperate Loliinae (Poaceae, Pooideae) grasses in the northern and southern hemispheres. Mol. Phylogen. Evol. 46: 932-957.
Moore, G., Macklin, J. & L. DeCesare. 2010.
A brief history of Asa Gray's Manual of Botany. Harvard Papers in Botany 15: 277-286.
Pyšek, P., Richardson, D.M., Rejmánek, M., Webster, G., Williamson, M. & J. Kirschner. 2004.
Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53: 131143.
Raven, P.H. 1977.
The California flora. Pp. 109-137 in: M.G. Barbour & J. Major (eds.) Terrestrial Vegetation of California. John Wiley & Sons, New York,
Rejmánek, M. 2007.
Book review on Weeds of California and Other Western States by J.M. DiTomaso & E.A. Healy. Madroño 54: 361-363.
Rejmánek, M. & J.M. Randall 1994.
Invasive alien plants in California: 1993 summary and comparison with other areas in North America. Madroño 41: 161-177.
Romanchenko, K., Peterson, P.M., Soreng, R.J., Garcia-Jacas, M. & A. Susanna. 2010.
Phylogenetics of Stipeae (Poaceae: Pooideae) based on plastid and nuclear DNA sequences. Pp. 511-537 in: Seberg, O., Petersen, G., Barfod, A.S. & J. Davis (eds.) Diversity, Phylogeny, and Evolution in Monocotyledons. Aarhus University Press, Denmark.

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