| BEN |
| BOTANICAL ELECTRONIC NEWS |
|---|
| ISSN 1188-603X |
| No. 303 February 26, 2003 | aceska@victoria.tc.ca | Victoria, B.C. |
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It is with great sadness that I report the death of William Louis Culberson, distinguished Lichenologist, prominent botanist, and treasured friend of us all. He fought cancer for several months and died in his sleep with his beloved wife, Chicita, at his side on February 8, in Duke University Hospital. He was 73.
Bill received his B.S. degree at the University of Cincinnati, influenced greatly by E. Lucy Braun and Margaret Fulford, who steered him into Lichenology. He studied at the Universite de Paris, where he received an M.S. (Diplômé d'études Supérieures) and a Ph.D from the University of Wisconsin, where he worked with John Thompson. Before coming to the Department of Botany at Duke University in 1955, he was a post- doctoral at Harvard University.
Bill had the good fortune to meet Chicita in a German class at the University of Cincinnati and they both were overwhelmed with each other. He persuaded her to come to Wisconsin and get an M.S. in chemistry while he pursued his Ph.D., which she did. This was the beginning of an incredible collaboration in lichen chemistry and taxonomy. It was more than a simple collaboration: it was a union of old-fashioned love and scientific talents. I don't think either could have had the success they produced without the other. They were a remarkable team in all respects.
I won't detail all of Bill's honors. There are many. He was president of the Botanical Society of America and the American Bryological and Lichenological Society and was editor of both journals of the two societies. He taught lichenology for many years at Duke and inspired students in many other fields. A note from Marshall Crosby is typical of the responses: "His lichen class was one of the best courses I took at Duke, and he helped tremendously with my developing interests in editing and bibliography." Bill was a linguist. Unlike most American scientists, he was fluent in French and German. He knew Latin and wrote all of our Latin diagnoses. I don't know what we will do without him.
What many of you might not know about Bill was his dedicated interest in all aspects of horticulture and gardens. He was director of the Sarah P. Duke Gardens for 20 years and greatly expanded and improved it, notably by the addition of a magnificent Asiatic garden, now named for him. He rescued it from an ordinary Kodachrome garden to a real Botanical Garden! In addition to these outside activities, Bill produced more than 100 scientific papers in lichenology and among other things, wrote a beautifully written article for each issue of Flora, the Garden's publication. It is hoped that these articles will be put together in a book.
Bill is survived by his wife, Chicita, and a family of devoted friends in the Department of Biology at Duke University. A commemoration will be held at the Sarah P. Duke Gardens at a date to be announced. Donations can be made to the Gardens or to the American Cancer Society.
[See also http://www.botany.hawaii.edu/cpsu/acharius/medalsp.htm - AC]
The identification of a naturalized grape in Ontario as Vitis labrusca x riparia and the resemblance of its offspring to the widespread native Riverbank Grape, Vitis riparia Michaux, suggests that: (1) some apparently native plants resembling V. riparia may be of hybrid origin, and (2) the relatively recent expansion of viticulture in parts of Canada to the north of traditional grape growing areas as a result of the availability of more recently developed cold-hardy vines may result in naturalization of unfamiliar grapes. These are not easily identified using currently available literature. A key to the Ontario wild grapes and cultivated is provided.
A wild grape found in eastern Ontario near Garreton by Mr. Bob Woolham (Ontario: Grenville Co., S. of Garreton, 7 Sept. 1999, P.M. Catling, 15 August 1999, B. Woolham, DAO), could not be assigned to any known taxon. It was growing with Riverbank Grape (Vitis riparia Michx.) and Red-Osier Dogwood (Cornus stolonifera Michx.) along a fence line in a place where no grape had ever been cultivated. The grapes were 1.2-2.2 cm in diameter and dark blue with a bloom. The relatively large size of the grapes suggested the Fox Grape (Vitis labrusca L.) but the vine had the arachnoid pubescence scattered in pale tufts, rather than brownish and completely concealing the leaf undersurface as in V. labrusca. The grapes were in clusters of 12-30 and were mostly 1.5-1.7 cm long. Tendrils were present in some cases at six successive nodes, but in some cases they were altogether absent for up to four consecutive nodes. The entire fruit slipped out of the rind easily when squeezed and the flesh was mostly white. No other comparable specimens were found in the Agriculture Canada collection (DAO) nor have similar specimens been seen during field studies of grapes throughout southern Ontario.
Bailey (1923, 1934) developed the concept of V. labruscana Bailey for the "Labruscan vineyard grapes" (including the cultivars Concord, Worden and Niagara). Those grapes are selections of V. labrusca or crosses of V. labrusca and V. vinifera L. Bailey noted that V. labruscana " retains the essential habit, foliage and flower characters of V. labrusca" and "shows more of the V. labrusca character than of any other." Since the Grenville County vine shows only a very scattered and pale arachnoid pubescence on the leaf undersurface, quite unlike the dense brown covering that completely obscures the leaf surface of V. labrusca, it is not "more nearly related to V. labrusca than to any other Vitis", and is consequently not appropriately placed with V. labruscana. In Bailey (1934) V. labruscana appears only once under "with a thick dense continuous indumentum that covers and obscures the leaf surface itself (not to be confused with leaves merely pubescent on ribs and veins or irregularly floccose or "webby" in which the interlacing threads tend to accumulate in shreds and tufts with leaf surface exposed)". Since the grapes had a loose skin readily separating from the flesh and were thus readily "pulped", the Grenville County vine is probably not closely related to V. vinifera and V. vinifera hybrids, especially those in which the latter predominates.
In recent literature (Moore 1991; Gleason and Cronquist 1991), the Grenville County vine keys to Vitis novae-angliae Fernald, considered to be the spontaneous hybrid of V. labrusca and V. riparia. Both occur in eastern Ontario, although the former is rare. Fernald described V. novae-angliae as a species on the basis of constant or more or less uniform plants in a distinct bottomland habitat in a number of locations in Maine. He referred to it as a "thoroughly fixed and constant vine." Admitting a possible "hybrid origin in the long distant past", he (Fernald 1917) described Vitis novae-angliae as a species on the basis of its constant characteristics, very distinctive flavor of the fruit. and occurrence beyond the range of the putative parents. He noted that it resembled V. labrusca but differed in the loss (with age) of arachnoid pubescence on the leaf undersurface. He noted that it differed from V. riparia in lacking the jagged-dentate leaves with prolonged teeth, the teeth of V. novae-angliae being "broad, scarcely prolonged deltoid." Here a problem is immediately apparent since the teeth of the Grenville County vine are prolonged and quite unlike the lectotype (Moore 1991) and syntypes of V. novae-angliae in GH, but similar to V. riparia. The latter is ruled out as an appropriate name for the Grenville County vine due to both the relatively large grapes and arachnoid pubescence on the leaf undersurface. The Grenville County vine is also unlike the type of V. novae-angliae in the fact that tufts of arachnoid pubescence are scattered over the leaf surface. It could be accommodated in a broad circumscription of V. novae-angliae, i.e. all forms of first and later generation hybrids involving V. labrusca and V. riparia, but does not match the type of the putatively constant taxon.
Thus it seems most appropriate to refer to the Grenville County vine as V. labrusca x riparia. It could have originated from either wild species or from a cultivated grape, possibly of complex hybrid origin. Over the past few decades new hardy grape hybrids derived largely from V. labrusca and V. riparia are being grown in parts of Ontario far to the north of the traditional grape growing areas of the Niagara Peninsula and extreme southwestern Ontario. These have a capability to escape from cultivation as shown by the fact that 8 of 15 seeds of cultivar Marechal Foch survived the winter of 1999-2000 outdoors at Ottawa and so did 7 of 14 seeds of cultivar St. Croix. These plants grew into robust vines (specimens in DAO) but did not flower.
A detailed comparison with the similar St. Croix cultivar suggested that the Grenville County vine was not referable to that taxon which lacks a medium density of prostrate hairs between the veins and has a more widely open sinus at the base of the leaf (P. Hemstad pers. comm. and notes on specimens at DAO). An escape may not look much like its cultivated parents due to a high degree of heterozygosity, but may also differ substantially from native species. The progeny of the Grenville County vine (9 of 15 seeds overwintered outdoors in Ottawa) were quite unlike the parent in having essentially glabrous leaves closely resembling those of V. riparia. They did not flower after 2 years in cultivation. These observations suggest that some of the apparent V. riparia growing without cultivation in Ontario may be of hybrid origin, and cannot be identified as such using key characters currently available.
The identification of the Grenville County vine draws attention to the fact that the classification of wild grapes on the southern Ontario landscape may be much more complex than is immediately apparent. Some apparently native plants resembling V. riparia may be of hybrid origin and naturalization of unfamiliar grapes may increase. These possibilities are of importance with regard to protection of wild grape germplasm as well as accurate characterization of native biodiversity.
Although it is unclear whether the Grenville County vine originated from cultivars of complex hybrid origin or from wild species, it clearly possesses characteristics of both V. labrusca and V. riparia and is thus appropriately treated as a hybrid of this parentage. A key to the species occurring in natural situations in Ontario follows. The extent to which V. vinifera is naturalized or persisting requires further study. The key is based largely on the work of Bailey (1934) and Moore (1991).
1. Mature leaves whitish (glaucous) beneath
....................................... V. aestivalis Michx.
1. Mature leaves green beneath.
2. Mature leaves arachnoid-pubescent beneath with a permanent
and continuous rusty felt, the lower leaf surface not
exposed with age.
3. Grapes in a short, simple cluster as broad as long
....................................... V. labrusca L.
3. Grapes in a large, shouldered, tapering, elongated
cluster
................................. V. labruscana Bailey
2. Mature leaves glabrous or with simple or arachnoid pubes-
cence scattered and especially on the veins, the lower
leaf surface well exposed with age (note that young and
unexpanded leaves may have a dense felty covering).
4. Skin and fruit closely adherent so that the rind does
not slip off
....................................... V. vinifera L.
4. Skin loose and separated from the flesh when ripe so
that the rind can be readily slipped and the fruit is
thus readily "pulped".
5. Leaves with prolonged lobes and teeth.
6. Leaf undersurface with arachnoid pubescence
scattered in pale tufts; grapes 15-17 mm long
....................... Vitis labrusca x riparia
6. Leaf undersurface lacking arachnoid pubescence,
the surface glabrous or with straight simple
hairs; grapes 8-12 mm long
........................... Vitis riparia Michx.
5. Leaves unlobed or with short, shoulder-like lobes
and relatively short and rounded teeth.
7. Expanding leaves densely felty (becoming entirely
glabrous with age); tendrils present at three to
many consecutive nodes; nodal diaphragms 0.3-1.1
mm thick; grapes 12-17 mm in diameter
........................ V. novae-angliae Fern.
7. Expanding leaves glabrous or sparsely arachnoid-
pubescent, not densely felty; tendrils or in-
florescences present at only two consecutive
nodes; nodal diaphragms 1-2.5 mm thick; grapes
(3)8-12 mm in diameter
.................................. V. vulpina L.
Mr. Bob Woolham of Garretton, Ontario and Dr. Helen Fisher of University of Guelph assisted in obtaining material. Dr. Peter Hempstead, Research Viticulturist at the University of Minnesota kindly examined material and provided extensive comments on its probable identity.
Henderson's checkermallow (Sidalcea hendersonii) is a gorgeous wildflower in the hollyhock family (Malvaceae) - one of the treasures of the Pacific Northwest coast. Evidence suggests that the plant has virtually disappeared from Oregon, although it was originally known from at least ten scattered locations in this state from the mouth of the Umpqua River in Douglas County to the Columbia River estuary and from there along the coast of Washington and British Columbia to Sayward, north of Campbell River on Vancouver Island. This is, or was, its worldwide distribution. Where it still exists, the plant is found in a unique habitat: places where fresh water from lakes or streams approaches the Pacific Ocean. Its typical locations are tidal river estuaries, estuarine islands, outlets of sand dune lakes, or where fresh water marshes drain to the sea. A map of its historical and present distribution is virtually one-dimensional stretching in discrete dots up the Pacific coast of northern Oregon and Washington, then becoming more scattered among the islands of Puget Sound and southern British Columbia.
Where did this unique and spectacular species evolve, how did it spread to these isolated locations, and why has it virtually disappeared from Oregon? These questions have not yet been answered, but it seems likely that the species moved primarily from south to north. At least this is my hypothesis based on the fact that the genus Sidalcea is more diverse in California than in other locations. The Jepson Manual lists 18 species of Sidalcea for California (some of these with numerous subspecies); floras of Oregon unusually recognize 13 species (several of these with subspecies); Hitchcock recognized 3 species in Washington (again with some subspecies), and British Columbia has only the single species, Sidalcea hendersonii.
In his 1957 monograph on the perennial members of the genus Sidalcea, C. L. Hitchcock concluded that the closest living relative of S. hendersonii was Sidalcea nelsoniana, another rare species known only from Oregon. Thus it seems possible that Henderson's checkermallow evolved in Oregon, adapting to its unique coastal wetland environment. How it subsequently spread along the coast is not known. I have heard no evidence that its seeds are especially adapted to withstand saline conditions. Perhaps whole plants with soil torn loose in storms rafted northward on currents, coming to rest in coves and estuaries.
The species is vulnerable due to various factors. The first set is due to the vicissitudes of the estuarine habitat. Here, where fresh water meets salt, habitats may fall prey to flooding, siltation, storm surges, battering by driftwood and long-term changes in sea level. Other dangers are the anthropogenic threats which arise due to land management such as forestry, urban or resort development, road, bridge and airport construction, and the introduction of weedy competitors. Research in BC has also shown that the plant’s gynodioecious reproductive mechanism makes it vulnerable when population size becomes low. Weevils are known to feed on the seeds, perhaps also threatening smaller populations.
Sidalcea hendersonii was first made known to science by plant explorers in Oregon. Pioneer botanist Louis F. Henderson receives credit for sending the type specimen to the Gray Herbarium where it was named by Sereno Watson in 1887. Henderson collected the specimen probably near present day Fort Stevens State Park on the Columbia River estuary in Clatsop County. Simultaneously, it was found by another Oregon pioneer, Thomas Jefferson Howell, near the mouth of the Umpqua River in Douglas County. It has not been seen in either location for many years. If Oregon is indeed the species’ site of evolutionary origin, as well as the site of its first discovery, it is sad to contemplate what has apparently happened to it here. The list (Part 2; will be in BEN # 305) shows 9 (or perhaps 10) historic sites in our state. Surveys will take place at these sites in 2003, but based on current local knowledge plus evidence from searches in the 1970s, Henderson's checkermallow has been found at only 2 sites in Oregon in the last 40 years. (At one of these sites, only a single stem was noted.)
Shockingly, for a species this rare and this vulnerable, Sidalcea hendersonii receives no active protection or regular surveys in Oregon or Washington at the present time. The situation is similar in British Columbia where sites are listed, but environmental protection is perfunctory. This remarkable species deserves to be closely studied, but this may not happen unless it is recognized as a species of concern by the agencies whose job it is to monitor and protect our rare plants.
I suggest that our aim as concerned botanists should be to urge that this species be moved up in status so as to receive both study and protection. I will petition the Nature Conservancy to give it a global rating of G3 ("Threatened or rare throughout its range"). I will also request that it receive formal listing by the Oregon Department of Agriculture. And I will petition the US Department of Fish and Wildlife to designate it a Federal Species of Concern.
I hope that all NPSO members and other concerned plant lovers and conservationists will join me in these efforts. I also hope that you will organize and take part in field efforts during the summer of 2003 to survey the 9 historic sites in Oregon, as well as other likely but as yet unexplored sites in estuaries and similar habitats. Henderson's checkermallow is a beautiful plant, a part of our natural heritage that is apparently slipping into extinction in our state -- it deserves our attention and our efforts!