Bryolog 34 (25 November 2023)


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

  • Registration is now open for SO BE FREE 28, 29 March–01 April 2024 at the Santa Rita Experimental Range in Green Valley, Arizona! MORE
  • CA Bryoflora Forum Zoom session 5 pm, Tuesday 5 December 2023. MORE
  • International Association of Bryologists virtual meeting, 13–14 December 2023, details forthcoming HERE
  • First ever chapter-wide monthly meeting on Zoom in January 2024! MORE
  • Sunday 04 February 2024, beginners’ moss and liverwort walk at Headwaters Forest Reserve Elk River Trailhead in Humboldt County, followed by a microscope session at College of the Redwoods. MORE
  • Bryophyte Workshop for the Shasta Chapter in Redding, 15–16 February 2024. MORE
  • Save the date for the “Introduction to Bryophytes” Jepson Workshop, 9–10 March 2024. MORE
  • Save the date for the ABLS meeting, 11–14 July 2024 at the Shawnee Lodge and Conference Center in West Portsmouth, Ohio, separate from Botany meetings.
  • Join the Bryophyte Chapter board as President Elect or Treasurer. MORE

Quarterly Report

  • Update on Torrey Pines bryophyte and lichen survey HERE

Timeless Bits

  • Notes on the Moss Family Bryaceae I, by John R. Spence HERE
  • Fall Photo Gallery HERE

CA Bryoflora Forum Zoom Sessions

The CA Bryoflora Forum renews its series of informal Zoom sessions for those interested in the bryoflora of Northern California with a discussion scheduled for 5 pm (Pacific Time) on Tuesday 5 December 2023. For those previously involved, you may use the old ID and passcode. If you don’t have it, please message me for an email response ( Subsequent sessions may be on a three or four week interval depending on participants and topics.

You are all recommended to follow the Zoom sessions to be soon inaugurated by the Bryophyte Chapter.

North Coast Walk and Scope Session

Sunday 04 February 2024 (rain postpones to Sunday 11 February 2024). Double billing: (1) a beginners’ moss & liverwort walk, then (2) a microscope session. The walk starts at 9:30 am at Headwaters Forest Reserve Elk River Trailhead in Humboldt County. A photoguide of the more spectacular bryophytes will be provided. We’ll be done walking around 11:30 am. People are then invited to move to College of the Redwoods in Eureka. At 1:00 pm we will reconvene in SCI 102 on campus to look through microscopes at organs of note in the lifecycle of the plants we saw on the walk. People can attend the morning and/or the afternoon session. Leaders are Paul Wilson and Karen Reiss. Register via Send questions to

Monthly Chapter Meetings on Zoom

We are excited to introduce online monthly chapter meetings beginning in January 2024! At each meeting we are planning to have a few short presentations by board members, liaisons, and chapter members, to share updates on their bryophyte projects. We will also have a Moss of the Month, where Jordan Collins will highlight a particular bryophyte, a Couplet of the Month, where Paul Wilson will help us to distinguish between two closely related species, sharing his screen to show key microscopic differences, plus plenty of time to ask questions, plan field trips and share our joy of these lovely little plants! If you would like to attend these meetings, please fill out our survey so that we can schedule a day and time that works for most interested parties. Time and date to be announced soon on our website and via email.

Shasta Bryophyte Workshop

Scot Loring will lead a bryophyte workshop for the Shasta Chapter, Thursday 15 February – Friday 16 February 2024. Thursday 15 February will be a PowerPoint presentation mixed with hands-on lab activities at Shasta College in Redding (room to be announced) from 1:00 pm to 4:00 pm. Friday 16 February will consist of field trips to local destinations beginning at 9:00 am, ending later in the afternoon. Check our Event Updates page or the Shasta Chapter website or email for updates on location details.

Introduction to Bryophytes

Brent Mishler and John McLaughlin will teach a Jepson Workshop on bryophytes, 9–10 March 2024 at the University of California, Berkeley, and Redwood Regional Park. This workshop will provide you with the opportunity to learn more about the biology and identification of mosses, liverworts, and hornworts. It will introduce you to the strange and wonderful world of these small but important plants. They are maximally different from their kin, the larger vascular plants; you’ll learn why it is said that “Mosses are from Mars, vascular plants are from Venus.” Plan to experience the 30th anniversary of this legendary workshop, first taught in 1994! All day Saturday, and Sunday morning, we will meet in the Valley Life Sciences Building (VLSB) on the UC Berkeley campus for presentations and a hands-on introduction to how to make microscopic preparations and identify bryophytes. Sunday afternoon we will migrate to a local field site for brown bag lunch plus an afternoon field trip to learn about these plants in the field.

For a preview, see recordings from the 2022 workshop.

Details and registration form will be available starting December 1st at:

Join the Board!

A volunteer organization like ours only runs if people step up and take part. Positions on the Chapter Board begin their two-year terms on 1 April, but terms are staggered such that two positions are up for election each year. This year, the President Elect and Treasurer positions will be voted on. The duties of the positions are spelled out here: If you are interested in running for one of these positions, please contact our current President, Ben Carter at by 15 December 2023. The election will be held in February 2024 and results will be announced in the next issue of Bryolog.

Lichen and Bryophyte Survey at Torrey Pines

Chris Wagner and her team of students have been conducting a thorough survey of lichens and bryophytes at the Torrey Pines State Natural Reserve in San Diego since February 2023. No bryophytes have previously been recorded from the site, and the team has now documented 52 species, including 36 mosses, 15 liverworts, and 1 hornwort. They have plenty more collections to identify, many of which are in the Bryaceae, and are planning a Bryum Keying Party in December. Among their most exciting finds are Epipterygium tozeri, Scleropodium californicum, Geheebia ferruginea, Fissidens curvatus, Gemmabryum brassicoides, Gymnostomum viridulum, Hennediella heimii, Hennediella stanfordensis, Pseudisothecium cristatum, and Tortula amplexa. For more info, contact Chris Wagner,

close up of moss
Gymnostomum viridulum


micrograph of leaf with gemmae
Gymnostomum viridulum leaf with gemmae


habit of moss
Geheebia ferruginea

Notes on the Moss Family Bryaceae I

— John R. Spence
California Academy of Sciences,

Welcome to the first in a series of Bryolog articles on the moss family Bryaceae. In this and future articles, I will explore topics including classification, species diversity, sexual and asexual reproductive biology, ecology and biogeography. This first installment looks briefly at different classifications of the family, and then examines species-level diversity, focusing on well-sampled areas including the Bryophyte Flora of North America region, the western Palearctic, and Australasia. This article also aims to establish a nomenclatural baseline of the family’s diversity for future installments.

The Bryaceae is one of the largest moss families, and is distributed worldwide from the Arctic to Antarctica, even occurring on nunataks within the Antarctic ice shelf. Although not ecologically dominant in most vegetation communities, the family is generally quite speciose regionally. Thus, it is hard to avoid the family unless one is determined to do so. Since many of the readers of Bryolog are in California and the west coast, I will use the California Bryaceae as a case study where relevant. Below I will briefly review various classifications of the family first, then examine world and regional level species diversity including the remarkable levels in California.

1. Classification

The Bryaceae is part of the order Bryales, which also includes several relatively advanced diplolepidous acrocarpous families. The classification of Brotherus (1924) was dominant throughout the 20th century up until ca. 2000, when molecular techniques were first developed and applied to taxonomy. There were many surprises of these early molecular studies. Brotherus recognized three subfamilies, all of which are now recognized as families: Bryaceae s.s., Mielichhoferiacae, and Orthodontiaceae. Pohlia and its allies were shown to be closer phylogenetically to the Mniaceae than to Bryaceae. Even more surprising, Leptobryum was moved to the Meesiaceae. Some rearrangements among the families were also made, for example species of Haplodontium being transferred to the Bryaceae s.s from the Mielichhoferiacae. Table 1 shows three current classifications, mine based primarily on my Australian and North American work (Spence 2014; Spence & Ramsay 2019), the recent European classification (Holyoak 2021), and the original Brotherus treatment.

Elsewhere I have discussed classification issues in the Bryaceae (Spence 2022). Some of the more recent work indicates that the genus Gemmabryum is not a natural group, as some of the tuber-forming species appear to be part of Imbribryum. There still remain several groups of taxa that are also not likely to be related to the core of Gemmabryum (which consists of the bulbil-forming species allied with G. dichotomum), but which have not been examined using molecular methods. Ptychostomum is another genus that clearly needs revision. Finally, unpublished data on several taxa of Plagiobryoides also indicate that that genus appears to be polyphyletic. All these results show that there is significant homoplasy within several of the larger gametophytically-defined genera, and that similar looking taxa are not necessarily related. For purposes of this discussion, however, I will continue to use my original classification as a heuristic device, with the caveat that some of the genera I have recognized will need revisions.

2. Species Diversity

The last attempt to examine both the classification and species diversity in the Bryaceae was by Ochi (1992). He recognized ca. 250 species worldwide, placed in four genera. Since then, it has become clear that his genus and species definitions in some cases were too broad. For example, he did not recognize most species that form tubers or bulbils, lumping them into a few broadly defined taxa. Anomobryum and Rhodobryum were placed into a broadly conceived Bryum. I have taken his results and developed a spreadsheet with all known recognized species, organized by genera, subgenera and sections (Table 2). In addition, I have included species richness for three well-studied regions, North America, western Europe and Australasia (Australia and New Zealand). The current world total is about 450 well-studied species plus ca. 120 more that have not been recently examined as to where they belong. I have assigned the first group to a specific genus, subgenus or section, although some of the sectional names are informal. Based on recent discoveries and the description of new species, I have also taken a stab at expected totals. This is highly speculative but may give some idea of the many taxa still to be either discovered or types examined. Also relevant is that in Ochi’s many monographs on the family, he listed numerous types that could not be located or in some cases were likely lost or destroyed, especially those of C. Mueller. This table should be considered a work in progress as changes are continually being made.

The totals for the three regions are ca. twice those of Ochi (1992). For example, he listed 44 species for Europe, while the total based on my species concepts is 91, admittedly much higher than the estimates in Holyoak (2021). Ochi listed 33 species for Australia and New Zealand, while my estimate is 63 species. Part of the differences relate to species definitions, with Ochi using very broadly defined genera and species, while I tend to be more of a splitter.  But new species have also been discovered in the last 30 years, and some monographic studies have accepted species that Ochi placed into synonymy. Like many other plant families, the Bryaceae consists of both large worldwide genera as well as restricted monotypic genera. The type for the Hawaiian species Mniobryoides degeneriae has not been located, while the rare Cape Verde Islands species Perssonia sanguinea has not yet been examined using molecular methods, but shows similarities to Haplodontium. Osculatia columbica is an old name for the species usually known as Brachymenium globosum, and is a unique widespread Neotropical species with highly unusual peristome morphology. The recent expansion of Osculatia to species of Gemmabryum and Imbibryum is highly problematic for several reasons, as it creates a polyphyletic taxon. First, Osculatia columbica is basal to a large clade based on molecular results, including six terminal taxa (Pedersen et al. 2006): Acidodontium, Anomobryum, Bryum, Gemmabryum, Haplodontium, and Imbribryum. The sole feature to justify this expansion is the presence of bulbils. Bryum has priority and also produces bulbils. In addition, bulbils are plesiomorphic for the family and are found elsewhere in the Bryales. In fact, four of the six terminal taxa in this clade (see Spence 2022) produce bulbils.

One of the most exciting things about the Bryaceae is that new species are being discovered on a regular basis. Even more surprising is that some of these new species have been found in well collected regions such as Europe and California. In the last few years, my colleagues and I have described and elevated several new species from California, with at least three additional taxa awaiting formal descriptions. There are also several distinctive individual collections which I have been unable to name. Intensive collecting in other less well-studied areas such as Chile and China have also recently yielded several taxa likely new to science. It seems probable that as future fieldwork is conducted in the world, numerous new species are likely to be discovered.

3. California as a case study

The Bryaceae of California is an interesting case study on speciation and diversity in the family. New finds as recently as 2022, such as an unknown Haplodontium and the first documentation of Plagiobryum zierii in the state, indicate that more discoveries are likely to be made. These finds bring the total for the state to 9 genera and 72 species, a remarkable level of diversity. Of these, 11 species (15%) are endemic or near-endemic to the state. The reasons for this will be discussed in future installments on ecology and biogeography of the family, but it is likely a combination of stable climates through the late Cenozoic, the complex geology and topography of the state, and the wide range of climates from arid through mediterranean to montane, coastal humid and alpine. Like other diverse moss families such as the Pottiaceae, Orthotrichaceae and Grimmiaceae, the Bryaceae parallels in many aspects the high levels of diversity and endemism in the vascular plant flora of the California Floristic Province.

In the next installment I will discuss the fascinating complexity of sexual reproduction in the family. Please feel free to email me with questions and suggestions.

4. References

Brotherus, V.F. 1924-1925. Musci (laubmoose). In Die Natürlichen Planzenfamilien, 2nd. ed., Vols. 10-11, ed. A. Engler. Leipzig: Wilhelm Engelmann.

Holyoak, D. T. 2021. European Bryaceae. viii + 344 pp. Pisces Publications, Newbury, Berkshire, U.K

Ochi, H. 1992. A revised infrageneric classification of the genus Bryum and related genera (Bryaceae, Musci). Bryobrothera 1: 231-244.

Pedersen N, Holyoak DT, Newton AE. 2006. Systematics and morphological evolution within the moss family Bryaceae: A comparison between parsimony and Bayesian methods for reconstruction of ancestral states. Molecular Systematics and Evolution 43: 891–907.

Spence, J.R. 2014. Bryaceae Schwägrichen, in Flora of North America EditorialCommittee (eds), Flora of North America North of Mexico, Vol. 28, Part 2. Oxford UniversityPress, New York, Oxford: 117-185.

Spence, J.R. 2022. Issues in Bryaceae taxonomy and potential future projects. The Bryological Times 154: 3-7.

Spence, J.R. & Ramsay, H.P. Revised keys and additions to the Australian Bryaceae (Bryopsida). Telopea 22: 99-134.

Table 1. Three generic classifications of the Bryaceae with all currently recognized generic names in the first column and their placement in the three systems (na=not applicable).

Genus Brotherus Holyoak Spence
Acidodontium Acidodontium na Acidodontium
Anomobryopsis Anomobryopsis na Anomobryum
Anomobryum Anomobryum Anomobryum Anomobryum
Brachymenium Brachymenium Brachymenium Brachymenium
Bryum Bryum Bryum Bryum
Gemmabryum Bryum Bryum/Imbribryum Gemmabryum
Haplodontium Haplodontium na Haplodontium
Imbribryum Bryum Imbribryum Imbribryum
Leptostomopsis Brachymenium na Leptostomopsis
Mniobryoides na na ?
Ochiobryum Bryum na Ochiobryum
Osculatia Brachymenium na Osculatia
Perssonia na na Haplodontium?
Plagiobryoides Bryum Ptychostomum Plagiobryoides
Plagiobryum Plagiobryum Ptychostomum Plagiobryum
Ptychostomum Bryum Ptychostomum Ptychostomum
Rhodobryum Rhodobryum Rhodobryum Rhodobryum
Rosulabryum Bryum Ptychostomum Rosulabryum
Genera 8 6 15

Table 2. Current species-level diversity including known species, likely number of expected species, and three regions that are well studied and sampled.

Genus Subgenus/Section Number of Species Expected Number BFNA Region Western Palearctic Australasia
Acidodontium 15 20 0 0 0
Anomobryum 26 30 3 5 1
Brachymenium Brachymenium 30 35 0 0 0
Orthocarpus 8 10 1 0 0
Bryum 37 40 10 5 3
Gemmabryum Gemmabryum 30 40 8 12 8
Tuberibryum 25 30 4 4 5
Caespitibryum 8 5 3 6 2
“Apiculata” 10 15 2 2 5
Haplodontium 11 15 4 2 0
Imbribryum 47 50 11 6 9
Leptostomopsis 8 10 2 0 1
Mniobryoides 1 ? 0 0 0
Ochiobryum 5 5 0 0 1
Osculatia 1 ? 0 0 0
Perssonia 1 ? 0 0 0
Plagiobryoides 20 25 5 2 2
Plagiobryum 9 10 4 3 1
Ptychostomum Cladodium 38 40 17 16 7
Ptychostomum 26 30 16 17 0
Rhodobryum 17 20 2 2 1
Rosulabryum Rosulabryum 45 50 4 1 8
Trichophora 30 35 11 8 7
Total Species 448 513 105 91 63
Unknown Affinities ~120 ? 3 0 0


wreath with mosses and fungi
Mossy log wreath, complete with fungi


River with trees along banks
Middle Fork Willamette River


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