ISSN 1188-603X

No. 533 March 5, 2019 Victoria, B.C.
Dr. A. Ceska, 1809 Penshurst, Victoria, BC, Canada V8N 2N6


From: Adolf Ceska

BEN is back! This time it is with a special bryological issue!

Last year, in 2018, I relaxed my BEN production effort and had serious thoughts about quitting BEN. I was deeply touched by the assisted suicide of my spiritual leader, botanist and plant ecologist David W. Goodall. My BEN effort started to seem meaningless. The work on this special, bryological issue of BEN was the last straw that broke the camel's back. "Why would I go into all those [editorial] troubles when nobody reads it anyway?"

After I announced the end of BEN (BEN has been posted since spring 1991), I was shocked. Within a few days, I received close to one hundred thank-you messages (97 or so, to be precise), some of them urging me not to quit. I have not realized that I had so many e-penfriends. I was forced to go back to the keyboard.

What more should I say? Thank you all who contacted me after my quitting message. Remember that BEN depends on you as contributors, authors of the tidbits suitable for posting. I would not have been able to post BEN without you, my contributors. I would especially like to thank Prof. Scott Russell, University of Oklahoma, who has been archiving BEN on the OU web pages since 1995. Thanks also go to all my close friends who have been translating my Czechlish into readable English. (I am now a good advertisement for the Grammarly with use of which my writing at least looks like English.)

Many thanks to all of you! Let's keep in touch!

DIANA HORTON (1949-2018)


Diana Horton died suddenly at her home in Vancouver, Washington, on June 24, 2018. Born in Edmonton, Alberta, Canada, Diana's parents, James Henry "Harry" and Betty Horton, instilled in her a life-long love of nature and the outdoors. Diana obtained a B.Ed. At the University of Alberta in 1972, she was followed by a Ph.D. in Botany in 1981. In 1983, Diana joined the Department of Botany at the University of Iowa. As an Assistant Professor, she became the Director and Curator of the Biology Herbarium at the University of Iowa, following in the footsteps of those responsible for the University's acquisition of specimens from around the world, since the 1870s.

Diana's specialty was the taxonomy, ecology, and phytogeography of bryophytes. In taxonomy she was a recognized expert on the genus Encalypta and she was active in phytogeography of bryophytes of NW Canada. Her knowledge and expertise were well-respected within the bryological community and will be missed. During her lifetime, Diana was active in many areas relating to the environment. A recipient of honours and awards, much of her time was spent teaching and interacting with her students, as well as with anyone who demonstrated an interest in knowing more about the natural world. Diana was, among other things, dedicated to the preservation of Iowa's native habitats through hands-on and collection-based research. Diana's tireless energy, on-going work in her fields of expertise, and her unflinching belief in the importance of fighting for environmental causes will be greatly missed.

An accomplished pianist, Diana enjoyed, joined, and supported a broad range of the Arts, such as opera, live theatre, music, and choral groups. During her lifetime, Diana gave a home to many rescue dogs. Her three most recent companions were with her when she died. Diana's death leaves a gap which cannot be filled in the lives of the close circle of her family, friends, and colleagues. A celebration of Diana's life will be held in Edmonton in the early summer of 2019.


From: Karen Golinski [] and Phil Burton []

The University of Northern British Columbia is offering an intensive field-based course in the biology of mosses and liverworts from April 29-May 9, 2019 at the scenic North Pacific Cannery National Historic Site ( ) at Port Edward, BC, near the mouth of the Skeena River, just south of Prince Rupert, BC. The 3-credit course is listed as BIOL 498 (Special Topics: Bryology) for undergraduates, or NRES 698 for graduate students. Graduate students from other universities in western Canada can apply for credit through the Western Deans' Agreement. Qualified individuals could also register for the single course under "interest only" or "audit" categories but must apply for admission to UNBC if not already a UNBC student.

For assistance with UNBC admission and registration, please feel free to e-mail or phone 250-615-5578.


From: Terry McIntosh

The 10th Annual Schofield Bryophyte Foray will take place in the south Okanagan Valley from May 30th to June 2nd. This year we will be exploring the dry interior of British Columbia, around Penticton with Terry McIntosh. We will visit open outcrops, shrub-steppe habitats, glaciolacustrine banks, as well as some dry Douglas-fir forest habitats. Discover numerous bryophyte species restricted to hot and dry areas of the province. Please contact Olivia Lee at for registration and more information. Also, it will be limited to 20 people.

[From May 30th to June 2nd, Pacific Northwest Key Council mycologists will have their spring foray at Silver Lake, Peachland, not far from Penticton. I am sure that the mycologists would appreciate, if bryologists would tell them about sites where they will see rich fruiting of mushrooms. AC]


From: Hawkins, B.J., May, E. & Robbins S. 2018. Nitrate and ammonium uptake in 21 common species of moss from Vancouver Island, British Columbia. Botany 96(3): 201-208. [Abstract] Contact:

Mosses play key ecological roles in water and nutrient retention in many ecosystems, yet relatively little is known of the functional characteristics of moss species, particularly nutritional characteristics. We investigated the net flux of ammonium, nitrate, and protons, using a microelectrode ion flux measurement system, in the gametophytes of 21 common species of moss from three contrasting locations in southern coastal British Columbia. The general location from which mosses were collected did not significantly affect ammonium or nitrate uptake. Proton efflux was greatest in mosses from locations with high rainfall. Rates of nitrate uptake differed among moss families, but there were no significant differences in uptake among species within families. Ammonium net flux differed among moss families, but also among species nested within family, with some species showing uptake and other showing ammonium efflux. In general, moss species native to dry habitats appeared to have higher rates of nitrogen uptake when ammonium and nitrate were available under favourable conditions.


From: Wu, E.T.Y., D.P. Mosquin, & R.D. Guy. 2018. An Inventory of Bryophytes on the Summit of Pink Mountain (Peace River District, British Columbia, Canada). Western North American Naturalist 78(1): 17-25.

Abstract The bryophyte diversity in northern British Columbia is insufficiently studied. We conducted a bryophyte survey in a 2.2-km2 proposed ecological reserve at the summit of Pink Mountain in the Peace River District of northern British Columbia. Pink Mountain is at the southern periphery of the ranges of several arctic flora species as well as the northernmost reaches of other temperate flora species. The south summit of Pink Mountain has a high floristic diversity coincident with its limestone geology, which provides a more alkaline condition for plant growth than the north summit does. We documented 65 species, including 2 new provincial records for British Columbia (Polytrichum hyperboreum and Tayloria hornschuchii), one red-listed (threatened) species (Tortula systylia), and one blue-listed (at-risk) species (Mnium arizonicum). Polytrichum hyperboreum and Tayloria hornschuchii have affinities to habitats more typical of polar regions. The presence of P. hyperboreum at Pink Mountain represents a southward range extension from 61°33' N in western North America. Because isolated populations at species distribution limits are likely to be vulnerable to climate change, the establishment of an ecological reserve on the south end of the summit of Pink Mountain is important for the preservation of bryophyte habitat.


From: T. McIntosh (, S. Joya (, and O. Lee (; UBC Herbarium, Botany Department, University of British Columbia, Vancouver BC.

Ongoing bryophyte inventory in British Columbia, coupled with literature reviews, have been rewarding over the past number of years, constantly adding to the biodiversity of the province. Twenty-two mosses are listed here. Of these, seven are reported new to both British Columbia and Canada and five new to B.C. Seven species have been previously reported for the province in earlier research papers but are not listed by the B.C. Conservation Data Center (CDC). Two species, Ptychostomum neodamense and Rosulabryum rubens, were reported in Volume 28 of the Flora of North America by John Spence, but herbarium specimens supporting their presence in B.C. are lacking at the Herbarium at the University of British Columbia (UBC). A recent email from Spence mentioned that he probably relied on literature reports for these occurrences, so we are listing them as new to the province. One species, Anoectangium sikkimense, will be reported as new to B.C. in an upcoming paper (Zander and Eckel. in prep.; with the authors' permission to include in this report). Eleven of the taxa are probable or possible introductions, usually restricted to urban habitats (collected mainly in Vancouver but also in Victoria by S. Joya).

Acaulon muticum (Hedwig) Müller Hal. var. muticum Distribution: new to B.C. (Mary Hill in Metchosin; Vancouver; Richmond), possibly introduced; Que.; Calif., Iowa, Kans., Mass., N.Y., N.C., Okla., Tenn., Tex.; Europe; Asia; Africa. Habitat: on small soil mounds in open ephemeral seep in Metchosin; on disturbed soil in Vancouver and Richmond. Representative Specimens: S. Joya 1422, 1427, 1428, 1953, 2080 (to be deposited). Notes: Acaulon muticum var. rufescens (A. Jaeger) H. A. Crum has previously been reported for B.C. (Priddle 1997). However, based on examination material from B.C. and elsewhere, there is some doubt whether varieties should be recognized (as done in the Flora of North America treatment; Zander 2007). The separation of the varieties, based on spore shape and ornamentation, which may best be attributed to phenological changes, as well as plant colour, appears spurious at best.

Acaulon triquetrum (Spruce) Müller Hal. Distribution: new to B.C. (Kamloops and northwest of Penticton); scattered across North America but mainly eastern: Ont., Sask.; Calif., Ill., Iowa, Mass., N.J., Tex., Va., W.Va.; Europe; Asia; n Africa; Australia. Habitat: on shaded, possibly seasonally damp silt, on the sides of glacio-lacustrine banks. Representative Specimen: T. McIntosh 9724 (UBC B229161). Notes: Both Acaulon species are tiny, appearing as small 'balls' on the soil surface, and can easily be missed by a casual observer.

Anoectangium sikkimense M.N. Aziz & Vohra Distribution: to be reported new to B.C. (Haida Gwaii, Toba Inlet, Skeena River area, Golden River), Canada, and North America in Zander and Eckel (in prep.); China, India. Habitat: humid cliffs and cliff crevices. Representative Specimens: W.B. Schofield 38343 (UBC B76064, originally identified as Anoectangium aestivum (Hedwig) Mitten), W.B. Schofield 21276 (UBC B75582, originally identified as A. aestivum (Hedwig) Mitten). Notes: The long-triangular and finely pointed leaves, as well as the usually serrulate basal leaf margins are characteristic of Anoectangium sikkimense. A review of herbaria specimens identified as A. aestivum will likely uncover more records of this and the following species for the province.

Anoectangium stracheyanum Mitten Distribution: reported as new to B.C. (Sims Creek Valley, Brooks Peninsula, North Vancouver, Revelstoke area, Haida Gwaii), Canada, and North America by Zander and Eckel (2017); China, Dominican Republic, India. Habitat: humid cliffs and cliff crevices. Representative Specimens: W.B. Schofield 67663 (NY, UBC B76077, originally identified as A. aestivum), W.B. Schofield 34006 (UBC B76057, originally identified as A. aestivum). Notes: The long-lanceolate to long-ligulate leaves that are often constricted just above the weakly sheathing leaf base are characteristic of Anoectangium stracheyanum.

Campylopus subulatus Schimper Distribution: new to B.C. (Lasqueti Island, Salt Spring Island, and the Greater Victoria area) and Canada; Calif., Oreg.; Europe; Asia. Habitat: in open sites on soil over rock, often with seepage. Representative Specimens: N. Djan-Chekar 2078 (UBC B150105), S. Joya, with A. MacKinnon, K. Luther, K. Brothers, and J. Ussery, 2148 (UBC B238875). Notes: A number of specimens of Campylopus subulatus are present in the Herbarium at UBC. However, it is not listed by CDC, probably because it usually has been lumped with C. schimperi Milde. Its subquadrate distal laminal cells and the lack of abaxial stereids in the costa help to identify the species. Campylopus subulatus is a southern, principally Mediterranean-type climate species.

Ephemerum serratum (Hedwig) Hampe Distribution: B.C. (Victoria, Vancouver; probably introduced), N.B., N.S., N.W.T., Ont., Que., Sask.; Ala., Calif., Conn., Fla., Ind., Ky., La., Maine, Md., Mass., Miss., Mo., N.H., N.J., N.Y., N.C., Ohio, Oreg., Pa., S.C., Tenn., Tex., Va., W.Va., Wis.; South America; Europe; Asia; South Africa; New Zealand. Habitat: mainly on disturbed soil, especially in bare patches in lawns on city boulevards, also on drying soil near pond. Representative Specimens: W. Miles 02-2001 (UBC B218648), S. Joya 1217 (UBC B210261), McIntosh 8002 (UBC B218647). Notes: Ephemerum serratum was previously been reported for B.C. (McIntosh and Miles 2005) but has not been listed by CDC. Species of Ephemerum are most commonly represented only as mats of filamentous green algae-like protonemata growing over bare soil. The production of tiny, rather narrow leaves sporadically occurs across these mats (at least in B.C.). Sporophytes, with ~1mm tall cleistocarpous capsules somewhat immersed in the leaves, are infrequently produced in B.C. The protonemata are branched and often more or less fan-shaped, in contrast to soil algal mats with less branched, disorganized strands, giving a clue to its identity when leaves are absent.

Ephemerum spinulosum Bruch & Schimper Distribution: new to B.C. (Kamloops; Salmon Arm); Ont., Que., Sask.; Ala., Ark., Conn., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Nebr., N.H., N.J., N.Y., N.C., Ohio, Pa., S.C., Tenn., Tex., Va., W.Va., Wis.; West Indies, Central America; South America; Europe; Asia. Habitat: on drying sandy-silts of eroding banks along Thompson River; on drying flats along receding lakeshore. Representative Specimen: T. McIntosh 9721 (UBC B218653). Notes: See comments under Ephemerum serratum.

Gemmabryum radiculosum (Bridel) J.R. Spence & H.P. Ramsay Distribution: new to B.C. (Vancouver) and Canada, probably introduced; Calif., Nev., Oreg.; Mexico; West Indies; Europe; Japan; north Africa; Canary Islands; Australia. Habitat: disturbed soil in gravel landscaping area along boulevard. Representative Specimen: S. Joya 124 (UBC B225405 as Bryum radiculosum Bridel; determined by J. Spence). Notes: Gemmabryum radiculosum is distinguished by the relatively large, reddish, spheric tubers and a long-excurrent costa.

Gemmabryum ruderale (Crundwell & Nyholm) J.R. Spence Distribution: new to B.C. (Richmond), probably introduced; Ont.; Ariz., Calif., La., Tex.; Eurasia; Africa; Macaronesia; New Zealand. Habitat: disturbed soil on boulevard. Representative Specimen: S. Joya 492 (UBC B225404 as Bryum ruderale Crundwell and Nyholm; verified by J. Spence). Notes: Gemmabryum ruderale is probably fairly common in the Vancouver area but underrepresented in collections. It is characterized by red-purple to violet rhizoids and spheric, purple-red rhizoidal tubers.

Grimmia moxleyi R. S. Williams in J.M. Holzinger Distribution: new to B.C. (Burnaby) and Canada, possibly introduced; Ariz., Calif., Nev.; Mexico. Habitat: on a wall on Simon Fraser University Campus. Representative Specimen: S. Joya 1467 (to be deposited; verified by Roxanne Hastings). Notes: The improbable occurrence of this southwestern species in an urban site in coastal B.C. is mysterious and may represent a rare introduction. The B.C. specimen is intermediate between Grimmia moxleyi and G. orbicularis Bruch, but has more characters in common with G. moxleyi. Neither species is previously known from Canada.

Hennediella stanfordensis (Steere) Blockeel Distribution: new to B.C. (Oak Bay, Victoria) and Canada, probably introduced; Calif.; Mexico; Europe; Australia. Habitat: bare, sometimes compact, disturbed soil. Representative Specimen: S. Joya 1644 (UBC B215574; verified by Richard Zander). Notes: Large patches of Hennediella stanfordensis observed in Oak Bay have likely been eliminated due to recent reconstruction of Oak Bay High School. All known observations to date are from within the vicinity of Bowker Creek, Victoria.

Imbribryum torenii J.R. Spence & Shevock Distribution: B.C. (Gulf Islands, Vancouver Island, and Vancouver); Calif., Ore., Wash. Habitat: sometimes seepy outcrops, mineral soil, gravel on roofs. Representative Specimens: K. Sadler 768 (UBC B2225594; determined by J. Spence), S. Joya, with O. Lee and A. Simon, 2025 (UBC B238521), T. McIntosh, with M. Fairbarns and M. Dunn, 10096 (UBC B239009; determined by S. Joya and T. McIntosh). Notes: This species was described by John Spence and Jim Shevock in 2015 (Spence and Shevock 2015). They listed it for B.C. but it is not listed in CDC. It is not rare along the coast, often overlooked or mistaken for similar species such as the common seepage site species Imbribryum miniatum (Lesquereux) J.R. Spence. Imbribryum miniatum has strongly concave leaves usually with rounded and cucullate apices whereas the leaves of Imbribryum torenii are only somewhat concave with acute, non-cucullate apices (also the costa is often short-excurrent in stout point, a feature lacking with I. miniatum).

Orthotrichum columbicum Mitten Distribution: B.C.; Calif., Oreg., Wash.; Europe. Habitat: mainly tree bark. Representative Specimen: S. Joya 2038 (UBC B238531). Notes: Orthotrichum columbicum has been, for the most part, long considered a synonym of O. consimile Mitten (e.g., Lawton 1971, Vitt 1973 and 2014) but was resurrected by Medina et al. (2012) who provided additional distinguishing characters and supporting molecular evidence. Medina et al. (2012) cite 19 collections of O. columbicum from B.C., indicating that it is fairly widespread along the coast. A more thorough review of specimens at UBC herbarium identified as O. consimile will undoubtedly uncover more records of O. columbicum for the province. A key to the species of the O. consimile complex is provided by Medina et al. (2012).

Orthotrichum cucullatum Lara, Medina, & Garilleti Distribution: B.C. (Vancouver), probably introduced; Calif. (possibly in Mex.). Habitat: tree bark. Representative Specimens: W.B. Schofield 87644 (UBC B117313; determined by R. Medina, originally identified as O. tenellum Bruch ex Bridel), S. Joya 1742 (UBC B225419; verified by R. Medina). Notes: This and the following species are in the Orthotrichum tenellum complex further clarified by Medina et al. (2013) following the description of Orthotrichum norrisii in 2008 (Medina et al. 2008). A key to these similar species can be found in Medina et al. (2013). All previous provincial records of Orthotrichum tenellum are referable to Orthotrichum cucullatum.

Orthotrichum norrisii Lara, Medina, Mazimpaka & Garilleti Distribution: new to B.C. (Vancouver) and Canada, probably introduced; Calif. Habitat: tree bark. Representative Specimen: S. Joya 616 (UBC B225413; verified by R. Medina). Notes: Plants from Vancouver differ from Orthotrichum norrisii in Medina et al. (2008) and Vitt (2014) in their apparent lack of a hairy vaginula that is said diagnostic of that species. Nevertheless, they otherwise agree with the concept of O. norrisii in the remaining characters, including the narrow, cylindric capsules with ribs that are only two cells wide.

Ptychostomum neodamense (Itzigsohn) J. R. Spence Distribution: new to B.C. (Vancouver), introduced in part, probably native elsewhere in B.C., Greenland; Alta., Nfld. and Labr., N.W.T., Nunavut, Ont., Yukon; Alaska, Calif., Colo.; Eurasia. Habitat: in gravel of landscaping material. Representative Specimen: S. Joya s.n. (UBC B225403; verified by J. Spence). Notes: Although it is listed as in B.C. by Spence (2014a), no specimens were located in the herbarium at UBC. Ptychostomum neodamense is closely related to two other B.C. mosses, P. bimum (Schreber) J. R. Spence and P. pseudotriquetrum (Hedwig) J. R. Spence & H. P. Ramsay ex Holyoak & N. Pedersen, differing in its ovate, blunt, mostly non-decurrent and crowded leaves. It may be native elsewhere in B.C.

Rosulabryum rubens (Mitten) J.R. Spence Distribution: new to B.C. (Vancouver; probably introduced), Que.; Calif., Md., N.J., N.Y., Okla., Tenn.; Europe; India; Australia. Habitat: on disturbed soil. Representative Specimen: S. Joya 546 (UBC B225406, as Bryum rubens Mitten; verified by J. Spence). Notes: As with Ptychostomum neodamense, although listed as in B.C. by Spence (2014b), no specimens were located in the herbarium at UBC. Rosulabryum rubens was probably introduced from Europe and is likely to be more widely distributed in North America, including other cities in B.C., than records show.

Scleropodium californicum (Lesquereux) Kindberg Distribution: new to B.C. (Greater Victoria area, including Trial Island and Sidney) and Canada; Calif., Ore.; Mexico (Baja California). Habitat: on rock and soil near shore. Representative Specimens: W.B. Schofield 77438 (UBC B117313; determined by S. Joya and verified by B. Carter; originally identified as Brachythecium starkei (Bridel) Schimper (= Sciurohypnum starkei (Bridel) Ignatov & Huttunen), S. Joya, with M. Fairbarns, 1987 (to be deposited). Notes: This species apparently has been overlooked, probably due to its superficial resemblance to other taxa, namely Homalothecium arenarium and Brachythecium albicans, which are found in similar habitats. Scleropodium californicum is more similar to species of Brachythecium, in particular B. albicans, but has distinctly thinner shoots and shorter laminal cells. From the other species of Scleropodium, it differs in being less julaceous when dry and growing in looser, thinner mats. Keys and illustrations for Scleropodium species can be found in Carter (2014).

Scleropodium julaceum Lawton Distribution: B.C. (Victoria and Saanich Peninsula), possibly introduced; Calif., Ore., Wash.,; Mexico (Baja California). Habitat: disturbed soil and bases of trees in mainly urban areas. Representative Specimens: F. Boas 215 (UBC B205252; determined by B. Carter, originally identified as Scleropodium cespitans (Müller Hal.) L. F. Koch., S. Joya 1634 (to be deposited). Notes: Scleropodium julaceum was first reported for B.C. by Carter (2014) based on a collection made by Frank Boas in 1962 near the Colquitz River at Beaver Lake, north of Victoria. Carter (2014) suggests that this species may be introduced in the northern part of its range.

Scleropodium occidentale B. E. Carter Distribution: B.C. (incompletely known, though apparently coastal; probably not rare); Calif., Nev., Ore., Wash. Habitat: Similar habitats as S. obtusifolium (rock and soil in wet places); more specimens are needed to better understand its habitat preferences in B.C. Representative Specimens: W.B. Schofield 31400 (UBC B117898; determined by B. Carter). O. Lee s.n. (UBC B22159; determined by B. Carter). Notes: Scleropodium occidentale was reported for B.C. by Carter (2014). It is morphologically and ecologically similar to S. obtusifolium (Mitten) Kindberg and often difficult to distinguish from it (the most diagnostic character appears to be robust costa that ends in a spine). A more thorough re-examination of herbarium specimens of S. obtusifolium will likely uncover more records of S. occidentale for the province.

Syntrichia papillosa (Wilson) Juratska Distribution: new to B.C. (known only from a single tree in East Vancouver), introduced; N.S., Ont.; Ariz., Calif., Colo., Conn., Ga., Ill., Maine, Mass., Mich., Mo., Nev., N.J., N.Mex., N.Y., N.C., Ohio, Pa., Tenn., Tex., Utah, Wash.; Mexico; South America; Europe; s Africa; Falkland Islands; New Zealand; Australia. Habitat: tree bark. Representative Specimen: S. Joya 602 (UBC B206196). Notes: The noticeably incurved leaves of Syntrichia papillosa have unipapillose cells, with the papillae on the abaxial leaf surface, a strongly papillose-serrate costa, and numerous small brown propagula, also on the adaxial leaf surface.

Triquetrella californica (Lesquereux) Grout Distribution: new to B.C. (Galiano Island) and Canada; Calif., Ore. Habitat: on soil over open outcrop flat above bluff. Representative Specimen: O. Lee 3253 (UBC B233200; determined by S. Joya)., S. Joya, with O. Lee and A. Simon, 2023 (to be deposited). Notes: This North American endemic was discovered by O. Lee on Galiano Island and represents a large range extension from its southerly location in Oregon. Triquetrella californica is distinctive in having a prostrate and mat-forming growth form, unlike most members of the Pottiaceae which grow as erect stems. Also, it has rounded-triangular stems, ovate leaves with folded apices, and tall, often branched papillae on the leaf surface. It is considered rare across its range but more occurrences in open, Garry oak-related habitats along the B.C. south coast are expected.


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Macroevolutionary evaluation methods extended and exemplified with Anoectangium (Pottiaceae, Bryophyta) in North America and the Himalayas

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