Bryolog 37 (28 August 2024)

Upcoming

As a general policy, check the website for event updates before you walk out the door!

• Fourth Thursday monthly meeting 26 September 2024 at 7 pm PT on Zoom. MORE
• Please consider running for Field Trip Director or Secretary. Don’t be bashful. Fill out form to self-nominate or email a current board member.
• Registration is open for SO BE FREE 29, 30 May–02 June 2025 in Humboldt County, CA. MORE
• Join the California Native Plant Society for just $30 – 40% off the regular price of $50! Be sure to choose the Bryophyte Chapter as one of your two chapters.

Quarterly Report

• Many thanks to the Hillside Gardeners of Montclair, Oakland, for another generous donation to the scholarship fund, and to Kiamara Ludwig, our former Treasurer, for her tireless fundraising efforts.
• Student research grant awarded to Zane Walker. MORE
• Peri Lee Pipkin, 2023 student research grant recipient, reports 16 bryophyte taxa new to Esmeralda County, NV. MORE
• Read about all of our student grant awardees and donate to the scholarship fund HERE.
• We enjoyed many fantastic talks at our monthly meetings. Recap HERE

Timeless Bits

• Notes on the Bryaceae II: Ecology, by John R. Spence HERE
• Studies of Conocephalum conicum in North America, by David H. Wagner HERE
• Bryophytes of Coyote Ridge and Flat, Inyo County, California, by Martin Purdy, our 2019 and 2021 Student Research Grant Awardee HERE

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Student Research Grant Recipients

We are pleased to announce Zane Walker as the recipient of our most recent student research grant. Zane recently graduated from Oregon State University Honors College with a BS in Botany, and is beginning a PhD program at the University of Kansas, where he will continue his work in paleobryology. Zane wowed us with a glimpse into the world of moss fossils at our monthly Zoom meeting in July. The chapter awarded him $1,000 to support his work completing a Bryoflora of the Marble Mountain Wilderness in Siskiyou County, CA. Congratulations, Zane!

Peri Lee Pipkin, our 2023 student research grant recipient, recently graduated with an MS in Botany from the California Botanic Garden. Our grant helped to fund their master’s thesis research, A Floristic Inventory of the Vascular and Nonvascular Plants of the Silver Peak Range in Esmeralda County, Nevada. They made 68 collections of bryophytes representing 26 minimum rank taxa across 14 families, including 16 county records new to Esmeralda County. Of note was Amblyodon dealbatus, a moss collected only a couple of times in the entire state of Nevada. See the list of new records for Esmeralda County below.

Family	Taxon
Meesiaceae	Amblyodon dealbatus
Brachytheciaceae	Brachythecisatrum fendleri
Pylaisiaceae	Buckia vaucheri
Ditrichaceae	Ceratodon purpureus
Encalyptaceae	Encalypta intermedia
Encalyptaceae	Encalypta rhaptocarpa
Encalyptaceae	Encalypta vulgaris
Grimmiaceae	Grimmia montana
Grimmiaceae	Grimmia pulvinata
Orthotrichaceae	Orthotrichum hallii
Bartramiaceae	Philonotis fontana
Pseudoleskeellaceae	Pseudoleskeella tectorum
Grimmiaceae	Schistidium heterophyllum
Pottiaceae	Syntrichia papillosissima
Pottiaceae	Syntrichia princeps
Pottiaceae	Syntrichia ruralis

Check out the complete list of student research grant recipients, with photos and descriptions, on our new Student Research Grants page. And be sure to hit that donation button and make a donation to fund future budding bryologists!

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Monthly Chapter Meeting Recap

If you haven’t been attending our monthly meetings on Zoom, here’s a taste of what you’ve been missing.

At our May 2024 meeting, Paul Wilson presented Today’s Taxon, Physcomitrium, including Californian representatives, as well as the model moss, P. patens.

Ben Carter illuminated the finer points of identifying the six species of Scleropodium, in the field and through the microscope, complete with handy charts, drawing on his Ph.D. dissertation research.

In June, Russ Kleinman and Karen Blisard started us off with Today’s Taxon, Takakia lepidozioides, a moss that looks like a liverwort and puzzled bryologists for decades. They had recently traveled to the Pacific Northwest in search of Takakia, and shared a flurry of fabulous photos, with labels and descriptions.  

Jordan Collins then brought us on a wonderful tour of the many habitats of the Utom/Santa Clara River Watershed in southern California, and their bryophyte inhabitants, sharing some of his recent work as a CNPS field botanist.

For July’s meeting, Jordan Collins introduced us to the leafy liverwort Chiloscyphus, acquainting us with its habitat, compatriots and helpful key characters.

Zane Walker then impressed us with his talk entitled, “Growing in Gondwana to Frozen Fossil: An Anatomically Preserved Dicrand Moss from Antarctica.” He shared his work on an exquisitely preserved fossil moss specimen from the Cretaceous, and filled us in with a brief summary of recent work from the growing field of paleobryology.

 In August, Jake Bauer took the reins for Today’s Taxon with a very informative introduction to Triquetrella californica, arousing excitement amongst us all to get into the field and search for this rare moss.  

Ben Carter and John McLaughlin followed up by sharing their recent work distinguishing between Anacolia baueri and A. menziesii using gametophytic characters in addition to the traditional (and rarely observed) sporophytic characters, inciting us to take a closer look at our collections.

A huge thank you to all of our speakers for making these meetings so informative, enriching and fun, and to our M.C., Jordan Collins, for his lively leadership, and to all of the participants for your discussion and support!

Want to give a talk? Email amandaheinrich777@gmail.com, jordancollins@cnps.org, or bryophyte@cnps.org. Meetings are held on the fourth Thursday of each month on Zoom at 7 pm PT.

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Notes on the Bryaceae II: Ecology

—John R. Spence
California Academy of Sciences, Bryum500.JS@gmail.com

In this second installment on the biology of the Bryaceae, I examine the ecology of the family, focusing on the species and habitats in the California region, but also briefly discussing habitats elsewhere in the world.

Ecology including habitat relationships of bryophytes is a large and complex topic. For a more detailed analysis see Janice Glime’s online ecology book which is an excellent introduction to the topic. I will provide a more general view on the habitat preferences of the Bryaceae, focusing on three basic aspects: light, water and substrate. Following this I will discuss broader vegetation community preferences for the California species in the family.

I will use some simple terms as generalized habitats including the following:

Aquatic: floating or submerged in water;
Saxicolous: found on various rock substrates, sometimes with thin soil layers;
Terricolous: associated with soil, sand, peat, mud, humus and gravel;
Epiphyte: growing on living trunks and branches of woody species;
Decayed Wood: growing on fallen logs, stumps, and other decayed wood.

Other aspects of bryophyte ecology include substrate factors, including chemical factors, especially whether the substrate is acidic, basic or neutral. Calciphiles are species that prefer or are sometimes restricted to strongly basic rocks and other substrates, while acidophiles prefer or are restricted to acidic rocks and substrates such as granite, various mafic rock types, acidic soils and humus in bogs, etc. However, many species can occur on a range of substrate chemistries and are generally described as neutral in preference. One additional useful term is rheophyte, where a species is associated with streams and rivers, either along the riverbanks or submerged. These species have the ability to survive flooding and scour that is a frequent factor in these habitats.

Most Bryaceae occur in open to partly open habitats, often disturbed. They can be characterized as mostly generalists, found in a wide variety of habitats except for dense forests, where they are rare due to heavy litterfall that can smother plants. One exception to this is Rhodobryum, which can be found on soil and mud in swamp forests and other forested areas where water is generally available. Some species are common on soil in agricultural and other disturbed habitats, including farms, grazed lands, soil banks, and roadsides. Species in Gemmabryum that produce bulbils and rhizoidal tubers are often common in these habitats (Porley 2008). The presence of specialized asexual structures likely confers advantages to species where routine soil disturbance occurs. Diversity in the family is often high in arctic-alpine regions, as well as rock outcrops and cliffs, roadcuts, soil banks, and riparian zones. In addition to low diversity in dense forests, the family is also rare in desert regions, with only a few species existing in these harsh environments. Epiphytes are common, especially species of Acidodontium and Brachymenium, in tropical montane forests, but rare in more temperate regions. Only two species can be regularly found on trees in North America: Leptostomopsis systylia and Rosulabryum laevifilum. These species typically occur on trunks of large old trees such as oaks and may extend to larger branches. Strictly aquatic species are not found in the Bryaceae, although several species can survive submerged in lakes and streams. For example, populations of Ptychostomum pseudotriquetrum can live at depths of >100 meters in clear Antarctic lakes, while Plagiobryoides vinosula can live submerged in perennial rivers, streams and springs if conditions are appropriate. One genus, Imbribryum, is characterized by species that occur at waterfalls and rock faces wetted by snowmelt and winter rains, and also rarely in hot springs. Species of Bryaceae are rarely dominant components of bryophyte vegetation. In a few cases, including in wetlands, and at waterfalls, wet rock faces and springs, they can be common and sometimes abundant. However, like many other smaller acrocarps, in more shaded and humid stable microsites pleurocarpous mosses usually dominate, often outcompeting and overtopping other bryophytes. Finally, an unusual and rare habitat occurs in the Arctic, where at least two species, including Rosulabryum rubens, colonize whale bones on beaches.

California vegetation is highly complex and variable across the state, and along with complex geology and topography, explains the rich diversity of the family. Deeply shaded forest sites such as redwood forests generally lack Bryaceae, with those present usually restricted to canopy gaps, exposed stumps, and vertical boulder faces. In more open woodlands, members of the family become more common, especially if there are a variety of different microsites, and Rosulabryum species are often the most common group. Grasslands and chaparral support several smaller species including Bryum and Gemmabryum that often produce rhizoidal tubers and leaf axil bulbils. Deserts generally lack Bryaceae, with only a few species present, especially Bryum argenteum var. muticum and Gemmabryum kunzei. A general rule of thumb is that higher cooler mountains tend to support more species compared with lowlands. This is especially true for Ptychostomum, in which the majority of species occur in wet to moist sites in montane, subalpine and alpine regions. Subalpine and alpine regions support many Bryaceae, including some of the state’s rarest species, such as Plagiobryum zieri, Ptychostomum arcticum, P. cernuum, P. nitidulum, P. schleicheri, Haplodontium macrocarpum, H. tehamense, and H. sp. aff. himalayanum. Riparian zones, due to their complex microhabitats and varying water levels support many species, including such distinctive species as Bryum flabelliformum in ed., Gemmabryum valparaisense, Plagiobryoides vinosula, P. pallescens, and P. pseudotriquetrum. Springs also support many of the same Bryaceae that are found in riparian zones. Finally, anthropogenic habitats support a few species, some of which are likely introduced. Bryum argenteum var. argenteum is ubiquitous, while a variety of small Bryum and Gemmabryum species that can cope with disturbance are often present. Some introduced species that occur in these areas include Gemmabryum eremaeum and G. demaretianum.

The Bryaceae of California consists of ~71 species in 9 genera, as defined in my treatment in the Bryophyte Flora of North America. Since then, there have been significant changes in classification based on molecular data (see my Bryolog article in issue 34). For this article, I have moved two species of Gemmabryum to Imbribryum (see list below) based on the most recent results. Also, an upcoming study has indicated that Plagiobryoides is polyphyletic, and a soon to be described species is not related to P. vinosula, instead it will be described in Bryum. The full list of species with their habitat preferences is found in the last section of this article. Four undescribed taxa that await formal publication are included in the list.

Below, general habitat preferences of each genus found in California are described, including general elevational zones (roughly low <3000’: middle 3000–7000’; high >7000’). These descriptions are based on my experiences collecting in the state, published studies and label information. However, the family continues to surprise us, as species often appear where they are not expected.

Anomobryum: exposed seepy soil and rock faces, often acidic, across a wide elevational range.

Bryum: terricolous or sometimes saxicolous, usually on moist to dry soil and rock, occasionally on wet soil across a wide elevational range.

Gemmabryum: a wide variety of habitats, with many found on disturbed moist soil, sometimes on rocks, usually in open habitats at low to moderate elevations.

Haplodontium: terricolous or saxicolous—seepy to damp soil banks or rock crevices at high elevations.

Imbribryum: predominantly saxicolous, often associated with seepy rock and waterfalls, often acidic and other mafic rocks, sometimes along streams, occasionally on damp soil in wetlands, across a wide elevational range.

Plagiobryoides: rheophytic species, generally associated with rocks in and near streams, occasionally in springs, mostly low elevations.

Plagiobryum: terricolous, associated with damp soil banks, patterned ground or snowbanks, subalpine-alpine.

Ptychostomum: variable in preferences, but most species are terricolous, on damp to wet soil in meadows, wetlands, springs and tundra, occasionally on decayed wood in forests, mostly middle to high elevations.

Rosulabryum: mostly terricolous or on decayed wood in a wide variety of habitats, occasionally also on rock faces, boulders and rarely serpentine outcrops, predominantly low to middle elevations.

Climate change is likely to have significant impacts on the Bryaceae as well as other bryophytes. In California the trend to increasingly variable rainfall, higher temperatures and widespread fires will likely impact many species. Most at risk are those associated with streams, springs and wetlands with potential declines in water availability in the future. Bryophytes are not adapted to fire, and widespread mortality generally occurs after hot crown fires, followed by colonization by a few generalists such as Bryum argenteum and Funaria hygrometrica and other early colonizers. One species, Gemmabryum vinosum, has shown the ability to survive cooler surface fires and resprout, but this is exceptional for the family (Spence and Kellman 2015). Higher temperatures are also going to affect exposed rock communities where heat loads may reach critical levels, killing saxicolous species. A recent review using native vascular plants in the California Floristic Province by Harrison et al. (2024) discusses some of the broader impacts of climate change. Similar analyses of how bryophytes will be impacted would be valuable, but more work is needed to develop the necessary geographic data to model bryophyte distributions across the state.

Below is a list of known California species of the Bryaceae and their main habitats, based on my experiences collecting in the state, published studies and label information. However, the family continues to surprise us, as species often appear where they are not expected. We still have much to learn about species’ habitat preferences in California. One useful study that analyzes ecological preferences for the family can be found in Hill et al. (2007) for the Great Britain/Ireland bryophyte flora. Two species, Ptychostomum cernuum and Rosulabryum bornholmense, have not been recently collected and may be either extinct or earlier reports may have been misidentifications.

Anomobryum

A. concinnatum: seepy acidic rock faces and soil in exposed sites.

A. julaceum: seepy rock faces and soil in exposed sites.

A. species A: an undescribed species from Lake County and surrounding areas found along streams, and on damp rock and soil, often in drier sites than the other two species. Abundant on Table Mountain on basalt.

Bryum

B. argenteum: widespread, found on soil and rock, often in disturbed sites.

var. argenteum: throughout the state, on disturbed sites, often nitrogenous.

var. muticum: harsh environments, especially in desert and alpine regions.

B. blindii: a poorly known terricolous species found on damp calcareous soil banks at middle to high elevations.

B. calobryoides: a species of waterfalls and wet soils at middle to high elevations.

B. chryseum: rare terricolous species, on dry soil or soil covered ledges in grasslands, open woodlands and chaparral at low elevations.

B. flabelliformum in ed.: rheophyte, on wet rocks and soil in steams and springs at low elevations. This species has previously gone under the name Plagiobryoides renauldii.

B. lanatum: terricolous or saxicolous, on dry soil or soil covered ledges and rocks in grasslands, open woodlands and chaparral at low elevations to middle elevations.

B. veronense: rare species found on damp soil and sandstone rocks at low to middle elevations.

Gemmabryum

G. badium: exposed dry soil and rock at middle to high elevations.

G. barnesii: terricolous, often on disturbed moist soil at low elevations.

G. brassicoides: saxicolous, on calcareous to neutral rocks, often sandstone, rarely on dry soil, at low elevations.

G. caespiticium: widespread on dry soil and soil over rock including disturbed sites, often in semiarid regions, at low to high elevations.

G. californicum: on exposed sandy soils and rock outcrops in open woodlands and grasslands at low elevations.

G. demaretianum: terricolous, on disturbed damp soil, oddly associated with US Forest Service stations, at low to middle elevations. Probably introduced.

G. dichotomum: terricolous, on damp to dry soil, often in disturbed sites, at low to middle elevations.

G. eremaeum: introduced Australian species found in botanic gardens along the central coast on disturbed soil.

G. gemmilucens: terricolous, on damp to dry soil, rarely on rock, often in disturbed sites, at low elevations.

G. gemmiferum: terricolous, on damp to dry soil, often in disturbed sites, at low elevations.

G. klinggraeffii: terricolous, on damp soil, often in disturbed sites at low elevations.

G. kunzei: on dry soil and rock in exposed sites, low to high elevations, common in drier arid and semiarid parts of the state.

G. radiculosum: terricolous, on damp calcareous soil, often in disturbed sites at low to middle elevations.

G. valparaisense: on calcareous wet rocks and soil over rocks along streams and at springs at low to middle elevations.

G. vinosum: widespread at low elevations on both soil and acidic rock, often in exposed sites, found on graywacke and other mafic rocks.

G. violaceum: terricolous, on damp soil often in disturbed sites at low elevations.

G. species A: terricolous, a rare local species found on damp soil in grasslands, related to G. dichotomum but with much larger bulbils and more distant stem leaves.

Haplodontium

H. macrocarpum: subalpine-alpine on seepy acidic soil banks and mineral rich rock.

H. tehamense: rare species on volcanic rock at high elevations on Mt. Lassen.

H. species A: a recently discovered species from subalpine n. California on wet soil and rock near snowbanks. It is similar to the Asian-Aleutian Islands species H. himalayanum.

Imbribryum

I. alpinum: on wet soil and mineral-rich acidic rocks at middle to high elevations.

I. gemmiparum: terricolous, on wet soil in calcareous springs at low to middle elevations.

I. microchaeton: on wet basic to neutral soil and rock from the lowlands to subalpine.

I. mildeanum: terricolous, wet acidic soil in wetlands at middle to high elevations.

I. miniatum: saxicolous, widespread at low to middle elevations on granite and other acidic rocks associated with waterfalls and spring snowmelt areas

I. muehlenbeckii: on damp to wet soil and mineral-rich acidic rocks at middle to high elevations.

I. subapiculatum: terricolous, on damp acidic soil, often in disturbed sites at low to middle elevations.

I. tenuisetum: terricolous on damp to dry soil, often disturbed, at low elevations.

I. torenii: predominantly saxicolous, widespread on basic to acidic rock and soil covered ledges in seepy sites from low to high elevations; often in drier sites than other Imbribryum species.

Plagiobryoides

P. vinosula: facultative rheophyte, often on rock or sometimes wet soil along streams and at springs at low elevations.

Plagiobryum

P. zieri: terricolous, on seepy soil banks and near snowbanks at high elevations.

Ptychostomum

P. arcticum: terricolous, on seepy soil banks and near snowbanks in the alpine.

P. bimum: terricolous or rarely saxicolous, on wet soil, soil covered ledges along streams, at springs and other seepy habitats, middle to high elevations.

P. cernuum: rare terricolous species, middle to high elevation wetlands.

P. creberrimum: widespread common terricolous species of woodlands and forests at middle to high elevations, sometimes on decayed wood or soil on rocks; in drier habitats than the related P. pallescens.

P. cyclophyllum: rare terricolous species, middle to high elevations wetlands.

P. inclinatum: terricolous and saxicolous, moist to dry calcareous soil and rock in exposed sites, middle to high elevations.

P. knowltonii: rare terricolous and saxicolous species, moist to dry calcareous soil and rock in exposed sites, alpine.

P. lonchocaulon: terricolous species of woodlands and forests at middle to high elevations; in drier habitats than the related P. pallescens.

P. neodamense: terricolous, calcareous wet soil at middle to high elevations wetlands.

P. nitidulum: rare terricolous species, on seepy soil banks and near snowbanks at high elevations; this species may be a variety of the boreal P. intermedium.

P. pacificum: on wet soil in middle to high elevations fens.

P. pallens: terricolous, middle to high elevations wetlands.

P. pallescens: an elevationally widespread common terricolous species on wet soil, often along streams, in wetlands and in seeps and springs.

P. pendulum: terricolous and saxicolous, moist to dry calcareous soil and rock in exposed sites, middle to high elevations. The correct name for this species may be P. compactum.

P. pseudotriquetrum: terricolous or rarely saxicolous, on wet soil, soil covered ledges along streams, at springs and other seepy habitats, low to high elevations.

P. schleicheri: a terricolous subalpine-alpine snowbank specialist on acidic soil.

P. turbinatum: terricolous, calcareous wet soil at middle to high elevations wetlands.

P. weigelii: terricolous, middle to high elevations wetlands.

Rosulabryum

R. bornholmense: terricolous, on damp acidic soil at low elevations.

R. canariense: predominantly terricolous, on dry to moist calcareous soil at low elevations.

R. capillare: on soil, soil over rock and decayed wood in shaded sites at low to middle elevations.

R. elegans: saxicolous species in calcareous rock crevices and soil-covered ledges in exposed sites at middle to high elevation. This species is atypical in Rosulabryum and has been recently moved to Ptychostomum.

R. erythroloma: a terricolous species also common on decayed wood. Common along the coast at low elevations; one of the few species found in redwood forests.

R. flaccidum: on soil, soil over rock and decayed wood in open to shaded sites at low to middle elevations.

R. sanguilentum: on soil, soil over rock and decayed wood in open to partially shaded sites at low elevations. This species has previously gone by the name R. gemmascens.

R. laevifilum: on soil, soil over rock and decayed wood and epiphytic on tree trunks in open to partially shaded sites at low elevations.

R. rubens: on disturbed calcareous soil and rock, often on concrete at low to middle elevations.

R. torquescens: predominantly terricolous, on dry to moist calcareous soil at low elevations, occasionally on decayed wood.

R. species A: a rare undescribed species on serpentine deposits, first located at The Cedars.

Literature

Harrison, S., J. Franklin, R.R. Hernandez, M. Ikagami, H.D. Safford, and J.H. Thorne. 2024. Climate change and California’s terrestrial biodiversity. PNAS Perspectives 121, No. 32 (https://doi.org/10.1073/pnas.2310074121).

Hill, M.O., Preston, C.D., Bosanquet, S.D.S. and Roy, D.B. 2007. BRYOATT. Attributes of British and Irish mosses, liverworts and hornworts. NERC Center for Ecology and Hydrology and Countryside Council for Wales. 88 pp.

Porley, R. 2008. Arable bryophytes. A field guide to the mosses, liverworts and hornworts of cultivated land in Britain and Ireland. WildGuides, UK.

Spence, J.R. and K. M. Kellman. 2015. New and interesting species of Gemmabryum J.R. Spence & H.P. Ramsay (Bryaceae, Bryopsida) from California and the West. Madroño 62: 124-135.

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Studies of Conocephalum in North America

—David H. Wagner
davidwagner@mac.com
August  16, 2024

There has been confusion about the taxonomy of Conocephalum conicum in North America since 2005, when Conocephalum salebrosum was described from European populations (Szweykowski, Buczkowska et Odrzykoski). Because of a confusing map in that publication, many have assumed that C. salebrosum was the best name for the plants found in North America. Clarity has come from more recent phylogenetic studies of rbcL sequences. The pioneering work by Miwa et al. (2009) showed that except for a single possible record of C. salebrosum in Colorado, neither true C. salebrosum nor C. conicum occur in North America. Instead, they identified two cryptic species, which they designated Type A and Type C, the former mostly in the east and the other in the west. It is possible that future work with anatomical characters of fresh plants will distinguish A type from C type. Akiyama, H. and Odrzyoski, I.J. (2020) suggest A and C could be treated as one species. The rarity of sporophyte production makes naming species on this basis not generally useful. Paton (2022) mentions that no fertile plants of Conocephalum had been found in the British Isles at the time she was writing.

I have begun a study of a feature of sterile plants that may help our understanding of Conocephalum diversity. Apparently there has never been a systematic examination of the ventral scale appendages found protecting the meristematic notch of dividing thalli. In the past few weeks I have isolated and measured these scale appendages of two specimens from Oregon and one each from Vermont and Tennessee. The size of the scale appendages appears to be quite consistent within a single clone, with the western plants having appendages nearly twice the size of the eastern plants. Hopefully, this will develop into a means of defining cryptic species. Here are plates of the four specimens that I studied. It’s a start!

 

 

 

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