Bryolog 23 (25 February 2021)


—Brent Mishler, Chapter President

  • SO BE FREE 25 newly revised dates: 30 April – 3 May 2021
    Due to the lag in COVID-19 immunizations, the date for SO BE FREE 25 has been moved back to 30 April – 3 May 2021, at Saratoga Springs in Lake County. We will follow all required social distancing measures, including masks. We will have all meals and gatherings outside, and will take all necessary precautions to keep everyone safe. So it will be a wonderful and safe event.I know we are all ready to get back out in the field, observe bryophytes, and interact with our fellow bryologists! All the information plus the registration form are here: There is a free SO BE FREE T-shirt for the first 25 registrants! As always we offer a student discount; this year we have added a payment plan for students. The new registration deadline is 1 April 2021.Even if you registered for the cancelled 2020 foray and let us hold your payment, you still need to register again for 2021 using the form—there is a place to indicate the amount from your 2020 payment you want to apply this year (or you can save your 2020 credit towards next year if you wish).It will be great to get together again. Please encourage students and others interested in learning bryophytes to attend. We will have a beginners’ session, as always, to bring people up to speed. See you there!
    Every year we elect two members of the Bryophyte Chapter’s Board of Directors. This year we are voting on Secretary and on Field-trip Director. We have one candidate for each position. Larke Reeber is the current Secretary and is running for her second term. Bill Thiessen is the current Field-trip Director and is running for his second term. Please vote HERE (historical link deleted)…

Quarterly Report

  • A virtual workshop was held Saturday 30 January 2021: “Wonders of a dryland moss: Syntrichia from genomes to ecosystems,” hosted in partnership between the Jepson Herbarium at UC Berkeley <> and the collaborative research project “Desiccation and Diversity in Dryland Mosses,” <> with funding provided by the US National Science Foundation under the Dimensions of Biodiversity program. Presentations were made by Matt Bowker, Kirsten Coe, Jenna Ekwealor, Kirsten Fisher, Javier Jáuregui-Lazo, Brent Mishler, and Mel Oliver on different aspects of the project, including genomics, population genetics, reproductive biology, biomechanics, physiology, systematics, and ecosystem function, interspersed with hands-on activities and discussion. There were 50 enthusiastic participants. In order to make the workshop available more broadly, recordings of all eight 15-minute presentations have been uploaded to YouTube
  • Otherwise, the quarter was disappointing, and we’re looking forward to the end of Covid days and Covid nights.

Timeless Bits

  • The genus Hypnum is no longer present in California, by Jim Shevock: MORE…
  • Repost of a comic bit on the biology of water bears (rated PG-13): LINK…
  • Species Spotlight: Frullania bolanderi and Frullania catalinae, by Amanda Heinrich: MORE…
  • David Wagner’s description and photos of Solenostoma schusterianum, newly discovered in California by Jim Shevock: LINK…

The genus Hypnum is no longer present in California!

—James R. Shevock, California Academy of Sciences

At the time of Hedwig, the pleurocarpous mosses were basically viewed as components of two major mega genera. One group centered on Neckera, and the other on Hypnum. During the 1800s, numerous subgenera and sections were created within Neckera and Hypnum to deal with the amazing diversity of the pleurocarps, especially as more collections arrived in European herbaria from other continents. Neckera and Hypnum were simply bursting at the seams. The diversity of ‘Hypnum‘ in the early years was mind boggling. In a way, the creation of more and more subsections or subgenera was the first step of splitting these two genera into more recognizable and evolutionary units. A quick Tropicos search of these two generic names will make it abundantly clear that both genera received a lot of names! Neckera received around 260 names for various sections and subgenera, and nearly 700 published species were described in Neckera. But, it was considerably greater for Hypnum, with nearly 400 names for sections and subgenera and over 4,000 species originally described as a Hypnum! Can you imagine the task of writing a key for nearly 4,000 Hypnum species?

As bryologists had an increasing number of specimens (especially from the tropics) arriving from worldwide travels and expeditions to examine and describe, more and more monographic studies were initiated during the late 1800s and continue to the present day. These studies caused many genera as well as families to be established, or genera to move about from one family to another. The names of many of the subgenera or sections of Hypnum eventually were elevated to genera, and several formed the basis for the naming of new moss families. Today, we can quickly see all of the moss genera and their family placements within the Hypnales that were spawned from the original concept of Hypnum and Neckera (Goffinet & Buck 2020). So every generation of botanists has to deal with new names for species they once knew by another name. Clearly, there are more name alterations yet to come.

Even with all of this activity during the past 150 years or so, there still remained a core number of species viewed as Hypnum. One of the great studies on this genus was initiated by Hisatsugu Ando (1922–2005). He started publishing on Hypnum in 1962 and he continued working on this group of mosses for the next 30 years. Ando became the leading authority on Hypnum and related genera, and many moss floras around the world used his species concepts. Ando recognized around 11 sections in Hypnum. In Crosby et al. (2000), Hypnum is viewed as containing 81 species. Everything else originally named as Hypnum has basically been placed in different genera. But work in Hypnum was not yet finished. With the advent of molecular data, additional hypotheses on the circumscription of Hypnum were to be tested. Recently, several phylogenetic studies have further evaluated many of the remaining species of Hypnum, and not surprisingly, other taxonomic placements and name changes have been proposed. One study (Câmara et al. 2018) focused on a portion of Hypnum within the concept of the moss family Pylaisiaceae proposed by Schimper back in 1860. Schlesak et al. (2018) dealt with some European taxa of Hypnum, and in this study, several new genera were established. Then the well-designed and detailed work by Kučera & Ignatov (2019) further expanded on the studies of Câmara et al. (2018) and Schlesak et al. (2018) by sequencing members representing all of the sections of Hypnum as recognized by Ando. Their results concluded that Hypnum, as viewed by Ando, still remained polyphyletic, and therefore, they placed additional Hypnum members into other genera and families. They established a new moss family, Jocheniaceae to accommodate Jochenia a genus of two species. This genus was first proposed in the Schlesak et al. (2018) work, but at that time they placed Jochenia provisionally within the Entodontaceae.

Now, with this little bit of historical background, we can discuss the species of Hypnum as viewed in Norris & Shevock (2004) and what has happened to them. In California, 6 species of Hypnum were recognized for the state bryoflora. These are: H. circinaleH. dieckiiH. lindbergiiH. revolutumH. subimponens and H. vaucheri. Now based on recent phylogenetic studies referenced above, none of these 6 taxa remain in Hypnum or within the Hypnaceae.

So here are the new names for these 6 species and the current family placement for them.

Hypnum circinale = Trochophyllohypnum circinale (Hook.) Jan Kučera & Ignatov [Pylaisiadelphaceae]. This is a monospecific genus.

The Pylaisiadelphaceae is likely a moss family not familiar to most California bryologists. This family was established by Goffinet & Buck (2004) and now contains 22 genera incorporating many genera that were traditionally placed within the Sematophyllaceae, a family dominated by subtropical taxa closely related to the Hypnaceae.

Hypnum dieckii = Calliergonellopsis dieckii (Renauld & Cardot) Jan Kučera & Ignatov [Pylaisiaceae]. This is a monospecific genus.

Hypnum lindbergii = Calliergonella lindbergii (Mitt.) Hedenäs [Pylaisiaceae]. This is a genus of about 8 species.

Hypnum revolutum = Roaldia revoluta (Mitt.) P.E.A.S. Câmara & M. Carv.-Silva [Pylaisiaceae]. This is a genus of 2 species.

Hypnum subimponens = Stereodon subimponens (Lesq.) Broth. [Stereodontaceae]. This is a rather large genus since many Hypnum species ended up in it.

Hypnum vaucheri = Buckia vaucheri (Lesq.) D. Rios, M.T. Gallego & J. Guerra [Pylaisiaceae]. This is a monospecific genus.

Some botanists might have the perception that name stability was more constant in the past, but every generation of botanists had to evaluate the arrival of new information. In what constituted a genus or a family were various proposed hypotheses based on gametophytic and sporophytic characters and the weight or importance given to each feature in such classification systems. Today we have the extra advantage to test these hypotheses with molecular data. While many of the genera and families envisioned by bryologists in the 1800s have been supported by phylogenetic studies, additional modifications to taxonomic placement and circumscription of taxa also are being proposed. The molecular data has created some ‘lumping’ along the way, but the clear trend is for more ‘splitting’, especially of large and diverse genera. So based on the increasing number of phylogenetic studies, we are seeing genera moved to other families, establishing new families, or resurrecting families long out of use.

Literature Cited

Câmara, P.E.A.S, M. Carvalho-Silva, D.K. Henriques, J. Guerra, S.R. Poveda & M. Stech. 2018. Pylaisiaceae Schimp. (Bryophyta) revisited. Journal of Bryology 40: 251–264.

Crosby, M.R., R.E. Magill, B. Allen & S. He. 2000. A checklist of the mosses. Missouri Botanical Garden, St. Louis.

Goffinet, B. & W.R. Buck. 2004. Systematics of the Bryophyta (mosses): from molecules to a revised classification. Monographs in Systematic Botany from the Missouri Botanical Garden 98: 205–239.

Goffinet, B. & W.R. Buck. 2020. Updated on-line classification of the mosses.

Kučera, J., O.I. Kuznetsova, Manukjanova & M.S. Ignatov. 2019. A phylogenetic revision of the genus Hypnum: towards completion. Taxon 68: 628–660.

Schlesak, S, L. Hedenäs, M. Nebel & D. Quandt. 2018. Cleaning a taxonomic dustbin: placing the European Hypnum species in a phylogenetic context. Bryophyte Diversity and Evolution 40: 37–54.

Species Spotlight: Frullania bolanderi Austin and Frullania catalinae A. evans

— Amanda Heinrich, Channel Islands Liaison

This summer I was thrilled to stumble upon a large patch of the leafy liverwort Frullania bolanderi growing on a leaning Coast Live Oak in a canyon. I was thrilled because (a) Santa Barbara County is not exactly filled with trees draped with leafy liverworts, and (b) it turns out Frullania bolanderi is easily identified in the field, so rewarding!

I began to hunt for more Frullania bolanderi, examining trees near creeks in rather deep sections of adjacent canyons. Lo and behold, I found several populations, including some which I had previously walked right by, failing to notice this tiny, delicate, dark reddish plant, clinging to the craggy bark. I usually found it in the company of at least smattering of Orthotrichum and what I initially mistook for Porella, but soon came to realize was Frullania catalinae, the only other species in the genus documented in mainland Southern California (F. californica occurs on the Channel Islands).

leafy liverwort with moss
1. Frullania bolanderi with Orthotrichum sp.

Frullania bolanderi ranges in color from green to dark reddish-brown. The main stems are rather tightly appressed to the substrate (typically the bark of angiosperms but occasionally rock and sometimes even other bryophytes or lichens) and from a distance may look like simply a discoloration of the bark. At closer range, the tiny stems (0.5–0.7 mm wide) spreading delicately from the central mass resemble the intricate patterns of a henna tattoo.

leafy liverwort with moss
2. Frullania bolanderi with Orthotrichum sp.

The leaves of Frullania bolanderi are complicate-bilobed, with two lobes of unequal size folded on top of one another. From above, one sees the dorsal lobes which are larger, rounded, and tend to overlap one another in a shingle-like fashion (incubous). From below, the smaller ventral lobes, or lobules, are seen more clearly. These lobules are said to be ‘galeate’ or ‘helmet-shaped’ and are basically little pockets presumably adapted to hold water, or conversely, when immersed in water on the microscope slide, to hold annoying air bubbles.

leafy liverwort as seen through a dissecting microscope
3. Ventral view of Frullania bolanderi showing galeate lobules. Air bubbles were removed by heating the slide with a match

The underleaves of F. bolanderi are also bilobed, or bifid, a feature which separates the entire genus from Porella, whose underleaves are not lobed. F. bolanderi underleaves often sport a tooth or two, and tufts of rhizoids.

close up
4. Frullania bolanderi bifid underleaves

The key distinguishing feature of Frullania bolanderi are the rather erect branchlets, on which the main leaves have detached (save a few funky ones at the very tip), leaving the stem looking quite naked. These wormlike projections are easily seen in the field with a hand lens.

leafy liverwort with narrow branches
5. Micrograph of Frullania bolanderi flagelliform branch tips

On closer inspection, most likely at home under the scope, one will see that these stems are covered with smaller modified underleaves (or sometimes more ‘regular’ underleaves), as well as what appear to be the stumps of the fallen leaves. No other species of Frullania (estimated to be 350–600 worldwide) displays these flagelliform branches with caducous (detachable) leaves which serve as asexual propagules.

microscopic view
6. Micrographs of Frullania bolanderi flagelliform branch tips showing underleaves and caducous leaf stumps

Apparently, some smaller populations of F. bolanderi may lack these distinctive naked branch tips, in which case the small shoot size (0.5–0.7 mm wide) along with the rather large helmet-shaped lobules can be used to separate the species from the four others in the genus in California. (For more detailed and professional descriptions see references below.)

Frullania catalinae has the distinction of being the only paroicous species in the genus in California, with perianths at the apex of a shoot and androecia adjacent on the same stem and on lateral branches. All other Frullania species in California (F. bolanderiF. franciscanaF. nisquallensisF. californica) are dioicous, with separate male and female plants. (In my limited experience thus far, I have found only male F. bolanderi plants and have failed to spot any females though they are reportedly around.)

liverwort with parts labeled
7. Photomicrograph of F. catalinae exhibiting paroicous condition with perianth and androecia on the same plant

The perianths of F. catalinae are pleated and have a narrow opening at the top, somewhat reminiscent of puckered fish lips, and often referred to as ‘beak-like.’ Early on, perianths contain archegonia, followed by a green immature sporophyte, and finally a mature sporophyte with a dark brown capsule, complete with spores and elaters. Ultimately the sporophytes emerge from the perianths, split along four lines of dehiscence and eject their spores with the help of spring-like elater cells.


capsule opening and then opened
8. Micrograph of dehisced F. catalinae sporophyte. Inset: mature but undehisced capsule dissected out and cut in half to reveal spores and elaters

8. Micrograph of dehisced F. catalinae sporophyte. Inset: mature but undehisced capsule dissected out and cut in half to reveal spores and elaters. Photos: Amanda Heinrich

The elaters can even be seen in the field with a hand lens. In the photo below, many of the capsules have burst, appearing rather hairy with their strung out elaters. One capsule in the center has yet to dehisce.

liverwort macrophoto
9. F. catalinae in the field with dehisced capsules

The androecia resemble brussels sprouts and contain several antheridia, which in turn contain the sperm.

male parts
10. In this micrograph the antheridia can be seen through the bracts of a F. catalinae androecium. Inset: An individual antheridium dissected from a F. catalinae androecium

F. catalinae also ranges in color from green to reddish-brown, presumably depending on the degree of sun exposure. Smaller patches of F. catalinae have the distinctive Frullania look, with tightly appressed branches stretching out elegantly upon the bark or rock. At 0.8–1.2 mm, these branches are still small but significantly larger than those of F. bolanderi.

leafy liverwort macro photo on bark
11. F. catalinae on Coast Live Oak

In larger mats of F. catalinae, the branches tend to pile up upon one another, which fooled me into thinking they were “just” Porella.

leafy liverwort
12. F. catalinae on Coast Live Oak

The complicate-bilobed leaves of F. catalinae are often squarrose, held out at a right angle from the stem. The ventral lobules are frequently not galeate, not helmet-shaped, but are explanate or flat, also reminscent of Porella. Oftentimes both explanate and galeate lobules can be found on the same branch.

I have frequently observed a single tooth on these explanate lobules though perhaps this is an anomaly as I did not read this in any published descriptions.

13. F. catalinae leaf with explanate lobule

As in all Frullania species, the underleaves of F. catalinae are bifid. This again is in contrast to the simple, entire underleaves of Porella. Many underleaves have tufts of rhizoids.

14. F. catalinae underleaf with rhizoid tuft

Frullania bolanderi is found throughout California and in Oregon, Washington, the Great Lakes region, and northeastern United States, as well as in Europe. Ken Kellman, Santa Cruz Liaison, reports that in Santa Cruz County F. bolanderi is found in drier locations such as open oak forests, compared to the locally more common species F. californica, which prefers more humid environments.

F. catalinae has been documented on the Channel Islands, as well as the adjacent coastal San Diego and Santa Barbara Counties, and as far north as San Luis Obispo and Monterey Counties. Ken reports that he has seen F. catalinae in Monterey County only on rock.

For more detailed and professional descriptions of Frullania bolanderiF. catalinae and other Frullania species, check out the references below.


Doyle, W. T., and Stotler, R. E., 2006. Contributions toward a bryoflora of California III. Keys and annotated species catalogue for liverworts and hornworts. Madroño 53: 152–154.

Atwood, J., 2017. Frullaniaceae. Bryophyte Flora of North America, Provisional Publication, Missouri Botanical Garden.

Hong, W. S., 1989. The genus Frullania in North America west of the hundredth meridian. The Bryologist 92: 363–367.

Lye, Kåre Arnstein, 2014. Ecology and distribution of Frullania bolanderi in Europe. Lindbergia, 37:6–21.

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