ISSN 1188-603X

No. 461 December 19, 2012 Victoria, B.C.
Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2


From: Judy and Geoff Godfrey -

It is with deep regret that we share the news that the distinguished hepaticologist Dr. Rudolf M. Schuster (Rudy) passed away on November 16, 2012, at the age of 91+1/2. Another of the botanical greats gone from this world.

It may surprise some botanists - as it did us some time ago - that Rudy's Ph.D. thesis was in entomology! For more details of his life, see the obituaries at these sites: and University of Massachusetts: ttps://

Through his books and papers on hepatics (liverworts), Rudy Schuster was a mentor for me (Judy) in my Ph.D. studies on hepatics at UBC, where Dr. Wilf Schofield was my advisor. Rudy's six-volume set of The Hepaticae and Anthocerotae of North America - East of the Hundredth Meridian was my hepatic bible, and is essential for botanists working with this group of plants in North America. I valued every detail in the descriptions of morphology, habitat, and associated species; in his often colourful footnotes; and in his fine illustrations. I shared what I was learning with Wilf Schofield. He told us much later that he had been waiting for a grad student to come along who wanted to study hepatics - so that he also could learn these plants at the same time.

This story of the liverwort Schofieldia is a bit of a side-bar, but it does explain how we came to know Rudy. During my early graduate student days with Wilf, Geoff had had a dream that we hiked into the mountains and found an emerald-green liverwort named "Tanzanium schofieldatum". A couple of weeks later, we did collect a gorgeous emerald-green liverwort in a subalpine meadow in the North Cascades Mountains. Looking at fresh plants, it was soon apparent that this was a very puzzling plant that didn't fit any description. I believed it was a new species and sent specimens to three well-known hepaticologists for a consultation. The first two responses (from overseas) said "not new", and gave me the names of two different known liverworts; I knew they were wrong. We opened the third letter from Professor Schuster with some trepidation, as we had heard he was a harsh critic who didn't suffer fools gladly. Rudy kindly confirmed my assessment, writing, "Congratulations!" He recommended that I do some additional work on the plant, as he suspected it would turn out to be a new, as yet undescribed genus - not just a new species. That proved true, and we had to tell Wilf that he, along with "Tanzanium's" species name from Geoff's dream, had to be "elevated". Thus "Tanzanium schofieldatum" (which Wilf had said was "poor Latin construct") became Schofieldia monticola, in honour of Wilf Schofield's contributions to bryology.

Since that time, Rudy collected the plant himself, studied it in more detail, and placed it into a special "basement group" with four other Gonwanalandic taxa from which the more advanced genera of the Cephalozioideae are derived. See:

Schuster, R. M. 1993.
On Cephalozia pachycaulis sp. nov. and the perimeters of Cephalozia. Bryologist 96: 619-625. Schuster, R. M. 1995.
Notes on nearctic Hepaticae. XX. On Schofieldia and evolution in the Cephalozioideae. Fragm. Flor. Geobot. 40: 39-46.

For many years, Schofieldia was known as endemic to the Pacific Northwest, occurring from Alaska to Oregon. More recently, however, it has been reported from Kamchatka, Russia.

That first correspondence on Schofieldia, and Rudy's continued support and advice, led to our friendship over the years. When we first met in person, Rudy and his first wife, Olga, invited us (along with our then 4-month old son) for dinner at their home in Massachusetts, near the University of Massachusetts at Amherst, and we enjoyed their well-known hospitality. Rudy barbecued steaks over the fire in their fireplace, and with wine, classical music, and good conversation, it all made for a memorable first visit.

Rudy's second wife, Marlene, brought us into more frequent contact again with Rudy over the last six years. During this time, we have been working to help him finish volumes 3 and 4 of his Austral Hepaticae. The first two volumes were published by Nova Hedwigia (J. Cramer, Berlin, publisher) in 2000 and 2002. Like the eastern North American hepatic volumes, these books are filled with the detail for which Rudy is famous (or infamous), and with hundreds of his highly valued, meticulously drawn illustrations. Rudy has told us that if he had to do it all over again, he would have started with the Southern Hemisphere hepatics, as that region is far more evolutionarily important and interesting.

Rudy always enjoyed a good discussion about interesting hepatic and systematic problems and questions, and we were always amazed at the breadth and scope of the experience and knowledge he could draw on from pure memory. We will miss these stimulating and enjoyable visits, his "You see my point!" punctuations, and his very presence.

For more information on Rudy Schuster and his work, we recommend the volume of papers prepared in 1988 in celebration of his life and work:

Engel, J. J. 1988.
Bryological Contributions Presented in Celebration of the Distinguished Scholarship of Rudolf M. Schuster. Beihefte zur Nova Hedwigia 90: 1-397. (J. J. Engel, editor, with Sinske Hattori, consulting editor).

The first paper in this compilation provides some biographical information and a list of Rudy's books and papers to 1988. We expect that this literature list will be updated in some future publication on Rudy's life work. See:

Engel, J. J. & A. Klekowski. 1988.
The life and travels of a twentieth century plant explorer; together with a comprehensive bibliography of RM Schuster's publications in bryology. Beihefte zur Nova Hedwigia 90: 1-15. (Amanda Klekowski, as a high school student, had interviewed Rudy to write a term paper on his biography.)

The authors of the above paper mention that, as of 1988, Rudy had described over 300 new species and over 70 new genera of hepatics, and there have been many more since then. (It makes us feel very humble!) Rudy described new taxa based on his own extensive fieldwork and subsequent meticulous study in his lab. He particularly emphasized the importance of examining hepatic specimens while they are still fresh, as oil-bodies (especially) and other characteristics are lost in dried plants. He had little or no patience with the practice of designating new taxa through the exercise of nomenclatural revision. More of Rudy's character comes through in other papers in this volume, particularly "Rudolph [sic] M. Schuster: The Early Years", by Lewis E. Anderson, then at the Department of Botany, Duke University.

Rudy conducted fieldwork in the Southern Hemisphere over four decades. A couple of his earlier papers convey his interest in hepatics and phytogeography in this part of the world:

Schuster, R. M. 1969.
Problems of Antipodal Distribution in Lower Land Plants. Taxon 18, No. 1, Smithsonian Summer Institute in Systematics 1968, Part 1, pp. 46-91.
Schuster, R. M. 1979.
On the persistence and dispersal of Transantarctic Hepaticae. Canadian Journal of Botany 57(20): 2179-2225.

We doubt that anyone will ever again be able to match what Rudy achieved in his many years of hepatic fieldwork around the world, the degree of detail in his studies and meticulous illustrations of hepatic specimens, and the volume of fine publications he produced. He still had so many ideas in his head for new topics to research and manuscripts to complete. We regret the loss to the botanical world of his decades of experience and knowledge, of a gifted intelligence and scholar. And we're sad to lose him as a colleague and friend.


From: Jennifer L. Modliszewski, Department of Biology, Duke University, Durham, NC e-mail:

Couplets 1-3 taken from Thompson (1993). Information in 4a, 4b, 5a, 5b, 6a, 6b compiled from Grant (1924), Pennell (1951) and Macnair (1989).

1a. Bracts at nodes subtending flowers completely fused around stem forming
    a more or less circular disk, glaucous......... M.glaucescens Greene 
1b. Bracts or leaves at nodes subtending flowers with petioles or fused
    around stem only at their bases, not forming circular disk, not glaucous
   2a. Leaves or at least some  pinnately lobed or dissected into narrow
       segments .................................... M. laciniatus A.Gray
   2b. Leaves  entire or  crenate, not pinnately lobed or dissected (but
       base often irregularly dissected or small-lobed)
      3a. Bract or leaf pairs at nodes subtending flowers linear to 
	    lanceolate,  not fused at base ... M. nudatus Curran ex Greene
      3b. Bract or leaf pairs at nodes subtending flowers ovate to cordate
          or round, sometimes fused at base around stem
         4a. Calyx much inflated, cup shaped and blunt (i.e. the upper calyx
             tooth is only slightly longer than	others .....................
                                                      M.platycalyx Pennell
         4b. Calyx somewhat inflated but not cup shaped, upper calyx tooth 
             1.5-3x longer than the others
            5a. Corolla 18-45 mm long, pistil exerted from the calyx, lower 
                calyx teeth straight in maturity
               6a. Corolla lobes smooth....... M. guttatus Fisch. Ex DC.
               6b. Corolla lobes having pointed tips ......................
                                                    M. cupriphilus Macnair
            5b. Corolla < 30 mm long, pistil scarcely or not exerted from
                the calyx, lower calyx teeth fold over sharply in maturity,
                nearly closing the orifice
               7a. Pistil included within or equal to calyx, corolla tube 
                   nearly cylindrical, plants from 5-50 cm tall, large ones 
                   with quadrangular winged stem, diploid ..................
                                                       M. nasutus Greene
	         7b. Pistil usually exserted from calyx (up to 3 mm) corolla
                   tube narrowly funnel-shaped (infundibular), plants 5-25
                   cm tall, stems tending to quadrangular but not winged,
                        M. sookensis B.G.Benedict, Modliszewski, et. al. 


Grant, A. L. (1924).
A monograph of the genus Mimulus. Annals of the Missouri Botanical Garden 11: 99-388.
Macnair, M.R. (1989).
A new species of Mimulus endemic to copper mines in California. Botanical Journal of the Linnean Society 100(1): 3-13.
Pennell, F. W. (1951).
Mimulus. Pp. 688-732 in Illustrated flora of the Pacific States. Vol. 3.
Thompson, D. M. (1993).
Mimulus. Pp. 1037-1051 in The Jepson manual, [1st edition]

See also:

Modliszewski, J. L. 2012.
Shy monkeyflower (Mimulus sookensis, Phrymaceae) has come out of hiding: new species of Mimulus for the Pacific Northwest and California. BEN # 460, October 16, 2012.


From: Rudolf Schmid(1) & Mena Schmid(2)
(1) Department of Integrative Biology, University of California, Berkeley, California 94720-3140, USA -
(2) Somerville, Massachusetts 02144, USA -

After the August 1974 fire in the upper Hall Canyon area on the southwestern flank of Black Mountain in the northwestern San Jacinto Mountains, Riverside Co., California, the United States Forest Service revegetated the burn in the mixed-conifer forest with the Sierra Nevada endemic Sequoiadendron giganteum (Cupressaceae). On 1 May 2009 a GPS census starting at the head of Hall Canyon revealed both in the canyon and upslope beyond it at least 157 individuals in the vicinity of the Black Mountain Trail, plus an outlier 450 m distant near the summit. This species alien to southern California is regenerating prolifically on Black Mountain, as revealed by multiple age classes, from juveniles (seedlings and saplings) about 20- 60 cm tall to young adult trees over 6 m tall, up to about 40 years old, and reproductively mature. The naturalized population (<7 ha in 2009) also appears to be spreading from its initial "small area"of introduction (<2 ha in 1974). Analysis of published print and Internet literature suggests similar post-fire naturalizations of Sequoiadendron giganteum in the San Gabriel and San Bernardino Mountains of Los Angeles and San Bernardino Counties. State and regional floras and checklists for California should acknowledge the naturalization of this species in montane southern California in the San Jacintos and possibly elsewhere.


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