Bryolog 33 (25 August 2023)


As a general policy, check Event Updates before you walk out the door!

  • Paul Wilson will lead a moss walk in the Arcata Community Forest on 10 December at 10:00 am. For updates, email
  • Save the date for the “Introduction to Bryophytes” Jepson Workshop, 9–10 March, 2024. MORE INFO
  • Save the date for SO BE FREE 28 (29 March to 01 April, 2024) at the Santa Rita Experimental Range, Green Valley, AZ. MORE INFO
  • Discounted CNPS memberships ($30) for new members this fall. MORE INFO

Quarterly Report

  • SO BE FREE 27 in the Sierra was beautiful. Photo report HERE, brief summary with additional photos HERE
  • New website! Check out We need better photos, especially ones that have a long horizontal theme with not much that’s important at the top or bottom, 1800 px wide by x 800 or more high. Drop photos in THIS FOLDER or email constructive criticism to
  • Paul Wilson gave a zoom lecture to the North San Joaquin Chapter.
  • Ben Carter became our liaison to the Bristlecone Chapter. John McLaughlin took over as liaison to the Santa Clara Valley Chapter. And Chris Wagner is now our official liaison to the Riverside-San Bernardino Chapter. Thanks, Ben, John and Chris!

Timeless Bits

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 given in December on:

SOBEFREE 28–Southern Arizona

river drainage surrounded by mountains

The upcoming 2024 SO BE FREE will be held from Friday to Monday, 29 March to 01 April, 2024, at the Santa Rita Experimental Range (SRER), near Green Valley, AZ. Located in the sky island region of Arizona within the Santa Rita Mountains south of Tucson, the SRER is embedded within a 52,000-acre experimental range, and is the longest continuously active rangeland research facility in the United States. The field station will allow opportunities to explore the bryoflora of the varied habitats to be found in the sky islands. The habitats in the immediate vicinity of SRER include mixed conifer forest, streams, grasslands, oak savannah, canyons, as well as Sonoran Desert. And don’t forget to bring your binoculars as this area is a birder’s paradise!

Accommodations at the SRER include two bunkhouse-style cabins with a communal bath house and communal kitchen. Camping is available onsite and at nearby campgrounds, and there are several hotels nearby in Madera Canyon and Green Valley, AZ. The Station also has a meeting room where we will enjoy meals, conversation, and some captivating evening presentations by our attendees. A separate lab facility with space for compound and dissecting microscopes will be available for use by our group. If you plan to stay off-site, consider making your reservations soon, as hotels may fill up with bird watchers at this time of year.

Stay tuned for registration details coming this fall.

cabin surrounding by flowering trees

Fall Discount for New Memberships

Have you always wanted to be a member of CNPS but never actually pulled the trigger? Here’s your chance to join at the discounted price of $30 (regularly $50). Only available through this link. Discounted price appears when membership is in cart. Be sure to choose the Bryophyte Chapter as one of your two Chapters!

SO BE FREE 27 Sagehen Creek Photo Report

With many thanks to our organizers—Fieldtrip Director Bill Thiessen, Treasurer Kiamara Ludwig, and our man on the ground Shane Hanofee.

This year we gathered for our annual foray in California’s gorgeous Sierra, just north of Truckee in the Tahoe National Forest, June 23–26, 2023.

We were based at the lovely Sagehen Creek Field Station, where we enjoyed our tasty meals catered by Mountain Magic in the presence of tall whispering trees.

cabin, food wagon, people

On Saturday, Brent Mishler led the traditional beginner’s walk around the preserve—education and entertainment wrapped into one.

people standing on trail

Other folks caravaned to Canyon Three Meadow along the Little Truckee River, where we encountered the likes of Riccia californica, R. cavernosa, R. sorocarpa and R. nigrella.

People standing in a grassland

After a brief respite, we continued on to the rushing Truckee River itself, at the Tahoe Pyramid Trailhead.

Three guys sitting on a log

In the evenings we gathered indoors for cozy microscope sessions where we poured over our collections and enjoyed lively conversation.

people sitting at a microscope table

On Sunday, many of us headed to the west side of the Sierra, where we explored the Loch Leven trail, just off of I-80.


The large boulders, sandy soils, seeps and pools hosted treats of many flavors, such as large Meiotrichum and Polytrichum species, gobs of Grimmia hamulosa, a variety of more petite Pohlia species than are common on the coast, and the tiny Cephaloziella divaricata.

person sitting on boulder

Codriophorus acicularis was a joy to discover, here photographed by John McLaughlin.

moss with sporophytes

John McLaughlin spotted this darling patch of Mannia gracilis.

complex thallose liverwort

One of the highlights of the trip was the fen located at Sagehen itself. After a meandering hunt through the forest we stumbled out onto the spongy cushion.

People in a fen

Colorful sundews were surrounded by a vast sea of Meesia triquetra and Drepanocladus aduncus.

sundue amid mosses

The experience was enhanced by removing shoes, squeezing the mossies between one’s toes, and becoming one with the fen.

foot in wet mosses

On the way home, several attendees stopped at Grass Lake Fen for more moss squishing fun. A good time was had by all, and we can’t wait to do it again next year.

person in large wetland

Read Brent’s archival brief with additional photos HERE.

A New Genus for a Special Californian Liverwort, Calasterella californica

—David G. Long, Royal Botanic Garden, Edinburgh, UK

Exactly 150 years ago, Coe Finch Austin (Austin 1873) distributed a series of dried specimens of North American Liverworts with printed labels under the title Hepaticae Boreali-Americanae Exsiccatae, of which number 153 was Fimbriaria californica Hampe ex Austin. This name had been coined but not validly published by G.E.L. Hampe, then validated by Austin in two lines of Latin on the printed exsiccate label: “Differt a F. lindenbergiana, Corda; pedunculo robustiore pallidiore fere nudo, perianthiis majoribus albidis, etc. HAB. California, Bolander, Bigelow.”

However, the genus Austin placed it in (Fimbraria Nees, sometimes spelled as Fimbriaria) turned out to be a synonym of the genus Asterella previously published by A.M.F.J. Palisot de Beauvois (1805). It was Lucien Underwood who correctly transferred the species to Asterella (Underwood 1895), as Asterella californica (Hampe ex Austin) Underw., where it has remained until now. Under this name, this robust and very distinctive member of the complex thalloid liverworts has become familiar to Californian bryologists as it is found throughout the length of California and only just into neighbouring states (Oregon, possibly Arizona, as well as Baja California Norte).

Until very recently, identification of the genus Asterella was made easy by the presence on the underside of its female receptacles (archegoniophores) of conspicuous cage-like pseudoperianths, each enclosing a capsule (sporophyte), allowing dispersal of its spores rather like the peristome of a moss capsule. Of the five genera in Aytoniaceae (Asterella, Cryptomitrium, Mannia, Plagiochasma and Reboulia) Asterella was the only one with pseudoperianths, making it unique in the family.

This situation (like that of some other old bryophyte genera) was blown apart by the advent of molecular systematics. As part of her doctoral study in Edinburgh (a monograph of the genus Mannia), Daniela Schill discovered that a long-accepted member of Asterella (A. gracilis) actually belonged in Mannia, even though it has a pseudoperianth. This conclusion, although discovered through DNA work, was corroborated by several morphological features (Schill et al. 2010) and the traditional definition of Asterella was undermined. Since then, molecular systematics has come on by leaps and bounds and its application to Asterella has led to segregation of the new genus Asterellopsis R.L.Zhu & You L. Xiang (Xiang et al. 2022) for the Sino-Himalayan species Asterellopsis grollei (D.G. Long) R.L.Zhu & You L. Xiang. The new segregation of Calasterella is the latest step in this process, and its isolation within Aytoniaceae has been indicated by several different molecular studies (Long et al. 2000, Schill et al. 2010, Villarreal et al. 2016) and by a more detailed phylogeny now under way (Long et al., unpublished). Calasterella has therefore now been published as a new genus (Long & Xiang 2023); its distinctive supporting morphological differences are outlined below.

My own studies on Asterella began following a suggestion by the late Riclef Grolle from Jena in Germany, that I take up his preliminary taxonomic work on the genus, to apply modern techniques such as scanning electron microscopy of spores and later molecular systematics. Revision of Asterella in Europe and Asia became the topic for my PhD study at Trinity College in Dublin, Ireland, and resulted in a published monograph (Long 2006). My earlier fieldwork in eastern Asia made this study possible. My broader interest in the genus worldwide was stimulated by this work, and I soon realized that one of the best hotspots for the genus was California and a planned collecting trip there became a reality in 1998.

Colleagues in California (Jim Shevock, Brent Mishler and Bill Doyle) helped me to plan this trip, and in March 1998 I embarked on a three week trip (disguised as a vacation) around California with my wife Siobhan and two young girls Kathleen and Nina in a frighteningly large RV, staying in a range of public campgrounds from the Coastal Redwoods in the north to San Diego in the south, literally a tour of known and some new Asterella localities. Our first encounter with Calasterella californica happened very quickly, just NW of Cloverdale in Sonoma County on 27 March. Over 18 days I collected it in 21 localities, in 12 counties ranging from Riverside County in the south, to Lake County in the north, at elevations ranging from 140 to 1190 m. The end of March and early April in California is the perfect season for collecting thalloid liverworts as many are growing vigorously and producing sporophytes.

In 2006, the late Bill Doyle of UC Santa Cruz sent me some excellent living specimens from California, including Calasterella. In 2017, 2018 and 2019, while on study visits to CAS, I was able to join SO BE FREE trips with Brent Mishler and Jim Shevock, on which Calasterella was seen again, including further north in Josephine County in Oregon, and more sites in California.

small child with liverworts
Fig. 1. Kathleen with some of our first Asterella collections on our 1998 trip—Calasterella californica (lower) and Asterella palmeri (upper), from Sycamore Creek, Fresno County.


person with camera at soil bank
Fig. 2. Habitat of Calasterella californica on roadside bank at Cataract Creek, Marin County, with Jian Liu.

The distinctive features of Calasterella, as summarized by Long & Zheng (2023) are the dioicous sexual condition, the strictly dichotomous branching of the thalli, the ventral scales with up to 4 subulate appendages, the receptacle almost always 4-lobed, borne on a stout naked stalk, the involucres beneath the receptacle deeply cleft, the spores lemon yellow in color with prominent trilete ridges, a conspicuous wing and without large areolae but with a fine mesh of alveolae on both surfaces. The other three Californian species now remaining in Asterella (A. bolanderi, A. palmeri and A. lindenbergiana) are all autoicous with male cushions borne on the same plants as the female receptacles, either on the main thallus just behind the base of the receptacle stalk, or on short branches arising ventrally.

complex thallose liverwort
Fig. 3. Calasterella californica, female plants with archegoniophores and pseudoperianths, at Cataract Creek, Marin County.


complex thallose liverwort
Fig. 4. Calasterella californica, male plants, from living collection supplied by the late Bill Doyle, from Pinnacles National Monument, San Benito County.


Much work remains to be done on Asterella and Calasterella in California, particularly on variation in spore ornamentation in Calasterella, where some spores show weakly to strongly prominent ridges as well as the fine reticulation. A study on Calasterella in California is currently underway at U.C. Berkeley by Ph.D. student Ixchel González-Ramirez, supervised by Brent Mishler, whose help is gratefully acknowledged, along with that of Jim Shevock and Dave Kavanaugh, California Academy of Sciences.

Literature cited

Austin, C.F. 1873. Hepaticae boreali-americanae. Published by the author, Closter, New Jersey.

Long, D.G. 2006. Revision of the genus Asterella P.Beauv. in Eurasia. Bryophytorum Bibliotheca 63: 1–299.

Long, D.G. & T.-X. Zheng 2023. A new subfamily Calasterelloideae and a new genus Calasterella for a phylogenetically and morphologically distinct member of Aytoniaceae. Phytotaxa 606: 225–230.

Long, D.G., Möller, M. & J. Preston. 2000. Phylogenetic relationships of Asterella (Aytoniaceae, Marchantiopsida) inferred from chloroplast DNA sequences. Bryologist 103: 625–644.

Palisot de Beauvois, A.M.F.J. 1805. Asterella. In: Cuvier, F. (Eds.), Dictionnaire des sciences naturelles dans lequel on traite méthodiquement des differents étres de la nature, fasc. 3. F. G. Levrault: Paris, pp. 257–258.

Schill, D.B., Long, D.G. & L.L. Forrest. 2010. A molecular phylogenetic study of Mannia (Marchantiophyta; Aytoniaceae) using chloroplast and nuclear markers. Bryologist 113: 164-179.

Underwood, L.M. 1895. Notes on our Hepaticae III. The distribution of North American Marchantiaceae. Botanical Gazette 20: 59–71.

Villarreal, J.C., Crandall-Stotler, B.J., Hollingsworth, M.L., Long, D.G. & L.L. Forrest. 2016. Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate. New Phytologist (2015) doi: 10.1111/nph.13716.

Xiang, Y.-L., Jin, X.-J., Shen, C., Cheng, X.-F., Shu, L. & R.-L. Zhu. 2022. New insights into the phylogeny of the complex thalloid liverworts (Marchantiopsida) based on chloroplast genomes. Cladistics 38: 649–662.


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