ISSN 1188-603X

No. 524 February 16, 2018 Victoria, B.C.
Dr. A. Ceska, 1809 Penshurst, Victoria, BC, Canada V8N 2N6


Wednesday, March 21, 2018
9:00 am - 4:00 pm
Reception 4:00 - 6:00pm

University of Washington Botanic Gardens
Center for Urban Horticulture, NHS Hall
3501 NE 41st St.
Seattle, WA 98105

An extensive network of professional, academic, and amateur botanists are actively engaged in the conservation, management, and study of Washington's diverse flora. Their expertise ranges from how best to manage biodiversity, to understanding climate change impacts on plant communities, to naming and classifying the flora's rare, common, and invasive elements. Invited speakers and poster presentations will share new insights and discoveries about these topics and more. Participants from throughout Washington and adjacent areas will have the opportunity to exchange ideas with colleagues within and across disciplines.
Co-hosted by the University of Washington Botanic Gardens and the University of Washington Herbarium at the Burke Museum.

Agenda, program information, and registration available at:

Student scholarships, remote attendance, and poster presentation options available. Visit website for details.

Questions? 206.685.8033


Bryce Kendrick. 2017. The Fifth Kingdom: An Introduction to Mycology. Fourth Edition. 514 p., ISBN 9781585104598 (paper), ISBN 9781585108626 (eBook) Hackett Publishing Company, Inc.

Now in full color and offering a wealth of new illustrations, the 4th edition of The Fifth Kingdom has been updated to reflect the most recent developments in mycology, including the field's adoption of a new taxonomical framework for fungi as a whole, and the latest advances in molecular genetics. The chapter on fungicides has been updated to include new discoveries. The discussion of poisonous mushrooms has been revised to include newly recognized types (and treatments) of mushroom poisoning. Chapters on medical aspects of mycology and practical uses for fungi have been expanded. Entirely new chapters-—on applications of mycological training, among other topics—-are all written with Kendrick's characteristic clarity, warmth, and humor—-the qualities that have helped establish The Fifth Kingdom as one of the best, and most engaging, introductions to mycology.

Bryce Kendrick has studied fungi for 55 years, during most of which he was a professor at the University of Waterloo. He has authored over 300 mycological publications, including several books. He was a Guggenheim Fellow in 1979, and was elected a Fellow of the Royal Society of Canada in 1981. He received the Distinguished Mycologist award of the Mycological Society of America in 1995, and was elected a Centenary Fellow of the British Mycological Society in 1996. In May 2001 he was the invited keynote speaker at the Mycological Society of Japan annual meeting in Tokyo, and in June 2001 received the Lawson Medal of the Canadian Botanical Association for lifetime contributions. As one of four authors, he has just helped complete a 1,000-page book The Genera of Hyphomycetes and a somewhat smaller book The Outer Spores: The Mushrooms of Haida Gwaii.


From: Garrett E. Crow, Prof. Emeritus, Univ. of New Hampshire

Utricularia ochroleuca R.W. Hartm., described in 1857, did not gain recognition as a species distinct from U. intermedia Hayne for some 57 years by European botanists. Since it is a species regarded as a vegetative apomict, purported of hybrid origin with U. intermedia and U. minor L. 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 and 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 A. Gray, 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 U. 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, U. stygia G. 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 shoots as well as the colorless shoots, 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; U. 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 U. 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 of 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, U. 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" U. 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, U. stygia, in North America.

In re-examining the specimens from the Gray Herbarium studied by Thor, as well as numerous additional specimens of the U. 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 U. occidentalis been determined conspecific with U. stygia, then the name U. 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. 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 some time ago I confirmed a second population (both non-flowering) discovered in Montana.


Ceska, A. & M. A. M. Bell. 1973.
Utricularia (Lentibulariaceae) in the Pacific Northwest. Madroño 22: 74–84.
Crow, G. E. 2014.
Lentibulariaceae [Pinguicula, Utricularia]. Pp. xx–xx, in: Flora of North America Editorial Committee. Flora of North America North of Mexico. Vol. 18. Oxford Press. New York, NY. Provisional Treatment:
Crow, G. E. & C. B. Hellquist. 2000.
Aquatic and Wetland Plants of Northeastern North America. Vol. 1. Pteridophytes, Gymnosperms, and Angiosperms: Dicotyledons. Univ. of Wisconsin Press. Madison, Wisconsin. i-lv + 480 p. 8 1/2 x 11, 338 line illus., 1 map
Plachno, B. J. & L. Adamec. 2007.
Differentiation of Utricularia ochroleuca and U. stygia populations in Trebon Basin, Czech Republic, on the basis of quadrifid glands. Carniv. Pl. Newslett. 36: 87–95.
Schlosser, E. 2003.
Utricularia stygia in California, USA, and U. ochroleuca at its southern range. Carniv. Pl. Newslett. 32: 113–121.
Taylor, P. 1964.
The Genus Utricularia L. Lentibulariaceae) in Africa (South of the Sahara) and Madagascar. Kew Bull. 18: 1–245.
Taylor, P. 1989.
The genus Utricularia - A taxonomic monograph. Kew Bull. Add. Ser. 14:1-724.
Thor, G. 1988.
The genus Utricularia in the Nordic countries with special emphasis on U. stygia and U. ochroleuca. Nord. J. Bot. 8: 219–395.

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