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Vol. 31, No. 3 July 2003 FREMONTIA

A JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

IN THIS ISSUE:

A CONVERSATION ABOUT MOSSES, LIVERWORTS, AND HORNWORTS by Dan Norris 5

MOSS GEOGRAPHY AND FLORISTICS IN CALIFORNIA by James R. Shevock 12 THE ROLE OF THE AMATEUR IN BRYOLOGY: TALES OF AN AMATEUR BRYOLOGIST by Kenneth Kellman 21

MOSSES IN THE DESERT? by Lloyd R. Stark 26 THE BIOLOGY OF BRYOPHYTES, WITH SPECIAL REFERENCE TO WATER by Brent D. Mishler 34 VOLUME 31:3, JULY 2003 FREMONTIA 1 SPECIAL ISSUE: BRYOPHYTES CALIFORNIA NATIVE PLANT SOCIETY FREMONTIA www.cnps.org MEMBERSHIP VOL. 31, NO. 3, JULY 2003 Dues include subscriptions to Fremontia and the Bulletin. Copyright © 2003 Mariposa Lily ...... $1,000 Supporting ...... $75 California Native Plant Society Benefactor ...... $500 Family, Group, International . . . $45 Patron ...... $250 Individual or Library ...... $35 Linda Ann Vorobik, Editor Plant Lover ...... $100 Student/Retired/Limited Income . $20 Daniel Norris & James R. Shevock, Convening Editors CONTACTS CHAPTER COUNCIL Bob Hass, Copy Editor CNPS, 2707 K Street, Suite 1 Alta Peak (Tulare) . . . . Joan Stewart Beth Hansen-Winter, Designer Sacramento, CA 95816-5113 Bristlecone (Inyo-Mono) ...... (916) 447-CNPS (2677) Stephen Ingram CALIFORNIA NATIVE Fax: (916) 447-2727 Channel Islands ...... Lynne Kada PLANT SOCIETY [email protected] Dorothy King Young (Mendocino/ Sonoma Coast) . . . . . Lori Hubbart Dedicated to the Preservation of Sacramento Office Staff: East Bay ...... Tony Morosco the California Native Flora Executive Director . . . . Pamela C. El Dorado ...... Amy Hoffman Muick, PhD Kern County . . . . . Laura Stockton The California Native Plant Society Los Angeles/Santa Monica Mtns . . . Development Director . . . Michael (CNPS) is an organization of lay- Betsey Landis Tomlinson persons and professionals united by an Marin County ...... Bob Soost interest in the native plants of Cali- Membership Coordinator . . Marin Milo Baker (Sonoma County) . . . . . fornia and is open to all. Its principal Lemieux Reny Parker aims are to preserve the native flora Sales Manager ...... Paul Maas Mojave Desert ...... Tim Thomas and add to the knowledge of members Bookkeeper . . . . Lois Cunningham Monterey Bay . Julie Anne Delgado and the public at large by monitoring Mount Lassen ...... Jim Bishop rare and endangered plants throughout Fremontia Editor . . . . . Linda Ann Napa Valley ...... Marcie Danner the state; by acting to save endangered Vorobik, PhD North Coast ...... Larry Levine North San Joaquin . . . . Bob Barzan areas through publicity, persuasion, Sr. Policy Analyst ...... Emily Orange County . . . Daniel Songster and on occasion, legal action; by pro- Roberson, PhD viding expert testimony to government Redbud (Grass Valley/Auburn) . . . . So. California Regional Botanist . . . Richard Hanes bodies; and by supporting financially Ileene Anderson Riverside/San Bernardino counties . . and otherwise the establishment of Rare Plant Botanist . . . Dave Tibor Katie Barrows native plant preserves. Much of this Vegetation Ecologist . . Julie Evens Sacramento Valley . . Diana Hickson work is done by volunteers through San Diego ...... Carrie Schneider CNPS Chapters throughout the state. Vegetation Ecologist . . Anne Klein San Bruno Mtn. Project Coordinator San Gabriel Mtns . . . . Lyn McAfee The Society’s educational work in- San Luis Obispo . . . . Dirk Walters Mary Petrilli cludes: publication of a quarterly jour- Sanhedrin (Ukiah) . Chuck Williams nal, Fremontia, and a quarterly Bulletin Legislative Advocate .Vern Goehring Santa Clara Valley . . Mary Simpson which gives news and announcements Legal Advisor . . . . Sandy McCoy Santa Cruz County . Janell Hillman of Society events and conservation Website Coordinator . Carol Witham Sequoia (Fresno) . . . . Marion Orvis issues. Chapters hold meetings, field Bulletin Editor . Michael Tomlinson Shasta ...... Dave DuBose trips, and plant and poster sales. Non- Sierra Foothills (Tuolumne, Cala- members are welcome to attend. BOARD OF DIRECTORS veras, Mariposa) . . . . Patrick Stone South Coast (Palos Verdes) ...... Money is provided through member Directors: dues and funds raised by chapter plant Barbara Sattler and poster sales. Additional donations, Carol Baird, Sue Britting, Steve Tahoe ...... Michael Hogan Hartman, Lori Hubbart, Lynne Kada, bequests, and memorial gifts from Willis L. Jepson (Solano) ...... Betsey Landis, David L. Magney, friends of the Society can assist greatly Mary Shaw Sandy McCoy, Patt McDaniel, J. Yerba Buena (San Francisco) . . . . . in carrying forward the work of the Spence McIntyre, Carol Witham Randy Zebell Society. Dues and donations are tax- deductible. MATERIALS FOR PUBLICATION Members and others are invited to submit material for publication in Fremontia. Fremontia logo (by L.A. Vorobik) reprinted Instructions for contributors can be found on the CNPS website, www.cnps.org, from The Jepson Manual, J. Hickman, or can be requested from Fremontia Editor, Linda Ann Vorobik, vorobik@rock Ed., 1993, with permission from the island.com, or c/o University and Jepson Herbaria, 1001 Valley Life Sciences Bldg. Jepson Herbarium, UC. © Regents of the #2465, University of California, Berkeley, CA 94720-2465. University of California. Prepress by ScanArt / Printed by Craftsman Trade Press

2 FREMONTIA VOLUME 31:3, JULY 2003 CONTENTS GUEST EDITORIAL ...... 4

A CONVERSATION ABOUT MOSSES, LIVERWORTS, AND HORNWORTS by Daniel Norris ...... 5 Through a series of frequently asked questions, Daniel Norris, who taught bryology for years, provides an introduction to the bryophytes (mosses, liverworts, and hornworts), as well as discussing some fascinating features of these small green plants.

MOSS GEOGRAPHY AND FLORISTICS IN CALIFORNIA by James R. Shevock ...... 12 Be amazed at the great diversity of mosses in California, the habitats in which they grow, and moss geography, along with a history of moss collecting in California, as you read this article written by one of California’s premier moss authorities. Then consider the fact that mosses new to California, including new species, are continually being discovered.

THE ROLE OF THE AMATEUR IN BRYOLOGY: TALES OF AN AMATEUR BRYOLOGIST by Kenneth Kellman ...... 21 Experience this journey of a commercial air conditioning mechanic with a fledgling interest 19 in mosses, to an amateur bryologist who is authoring a “Catalog of the Mosses of Santa Cruz County”! Ken Kellman shares his joy of learning more about mosses, along with tools, resources, and encouragement for others who may wish to travel this same road.

MOSSES IN THE DESERT? by Lloyd R. Stark ...... 26 Most of us think of mosses as being only abundant in wet habitats, festooning trees, or carpeting rocks and logs. In this article the author transforms these stereotypical images by writing of the many mosses of California deserts and where one finds them, along with many fascinating reproductive strategies employed by mosses surviving in such extreme habitats.

THE BIOLOGY OF BRYOPHYTES, WITH SPECIAL REFERENCE TO WATER by Brent D. Mishler ...... 34 Brent Mishler discusses the evolutionary history of bryophytes, and the physiological aspects of these plants that define where and how they live, with particular emphasis on mosses and water.

FROM THE ARCHIVES ...... 39

NOTES AND COMMENTS ...... 40

BOOK REVIEWS ...... 41

THE COVER: Gametophytes (green haploid generation) of Polytrichum commune, haircap moss. Photograph by M. Hutten. All images in this issue by M. Hutten unless credited otherwise. See this issue in color at www.cnps.org.

VOLUME 31:3, JULY 2003 FREMONTIA 3 GUEST EDITORIAL USEFUL WEBSITES AND CONTACT Ihave come to increasingly appreciate the abundance of opportunity to ex- INFORMATION plore and learn from the natural world. Why do I invest so much time and Bryophyte websites: energy in the study of plants? The answer is quite simple: it is fascinating. I learn See “Notes and Comments,” p. 34 something new almost every day. I can’t wait for my next hike where I might California Native Plant make an acquaintance with a new flower, and now, with each new bryophyte. Society: Botanical exploration has been both a journey toward intellectual enrichment as www.cnps.org, with links to conservation issues, chapters, well as my “mental health care package.” While our lives continue to become publications, policy, etc. more complex in our high-tech environment, we still have the opportunity to To sign up for “NPCC News,” e-mail news on native plant study nature and be amazed at the infinite ways in which it works. Several col- science and conservation, send leagues have asked me why I became interested in learning about the bryophytes. a request to [email protected]. The answer is not readily pinpointed, but as the following articles attempt to For updates on conservation convey, the bryophytes are, simply stated, a remarkable group of land plants. issues: They are worthy of our attention. Audubon Society www.audubon.org When we consider the rate of habitat conversion caused by various land-use Center for Biological Diversity www.sw-center.org practices, along with the growing population that continually impacts the Cali- Natural Resources Defense fornia landscape (e.g., urban sprawl), it is more important now than ever that we Council www.nrdc.org have conservation policies that provide for a network of protected areas. The Sierra Club protected places can (and should) run the full spectrum—from open space in www.sierraclub.org urban landscapes to the most remote areas in designated national wilderness Wilderness Society and national parks. All are important and contribute in their own way to species www.wilderness.org and habitat conservation. We are truly lucky to live in such a diverse and re- For voting information: markable state, and equally fortunate to have no less than 40 percent of our state League of Women Voters www.lwv.org, includes online voter in public lands. Few states can boast of such biological diversity as found in guide with state-specific nonparti- California. However, maintaining this heritage of biodiversity requires aware- san election and candidate information. ness, a desire to protect it, and a framework to actually manage for it. The be- US Senate ginning of this process is meaningful and long-lasting change, primarily through www.senate.gov education. Fremontia is one of these educational tools. US House of Representatives As much as we know about the California bryoflora, the process of gather- www.house.gov California State Senate ing inventory is still at an early stage compared with that for vascular plants. www.sen.ca.gov Many regions of the state are poorly collected, and our gathering of data about California State Assembly rare and threatened bryophytes and their habitats is still in the preliminary stages. www.assembly.ca.gov Nonetheless, we know that California harbors one of the most exciting and To write letters: species-rich bryofloras in the entire country. We are still discovering bryophytes President George W. Bush The White House that are new to science—currently we have more than two dozen in the process 1600 Pennsylvania Ave. NW of being described. This indeed inspires us to continue our exploration. I hope Washington, DC 20500 that through this bryophyte issue of Fremontia you may discover a sampling of Senator Barbara Boxer or Senator Diane Feinstein what motivates bryologists like me to spend days, months, and years with these US Senate little green plants. Washington, DC 20510 Your CA Representative —Jim Shevock US House of Representatives Bristlecone Chapter Washington, DC 20515 Member of the Fremontia Editorial Advisory Board

4 FREMONTIA VOLUME 31:3, JULY 2003 The moss Mnium spinulosum with gametophyte (leafy lower portion) and sporophyte (stalks and capsules). [See moss life cycle on page 28.] Note serrated margins toward leaf apex and strong costa (leaf midrib). A CONVERSATION ABOUT MOSSES, LIVERWORTS, AND HORNWORTS by Daniel Norris

any millennia ago, as a Bryophyta (or bryophytes) is the stem differentiation but instead MMichigan college fresh- scientific name for a group of plants grow as ribbon-like thalli (singular, man already ten years including the mosses, liverworts, and thallus). Bryologists typically study into bird-watching, I heard my hornworts. For our purpose, we can all groups of bryophytes despite the botany professor speaking of the define them as green plants without fact that the three groups probably joys of plant collecting. Why waste flowers and fruits, and lacking a well evolved independently from one time studying plants when beauti- defined system of vascular tissue for another. To avoid the more cum- ful birds beckoned? Two years later transporting plant fluids through- bersome term, bryophytes, I normally in Montana, I watched my pro- out the plant. They reproduce, not speak of mosses while meaning fessor’s wife collecting mosses while by seeds, but by single-celled spores. mosses, and liverworts and horn- I enjoyed the true excitement of Mosses and most liverworts have worts as a collective group. collecting beautiful flowers. Now, clearly recognizable leaves on clearly Over the course of my 45 years after 106,000 bryophyte collections, recognizable stems but they totally as a bryologist, I have gradually I continue to be obsessed with the lack a root system. All hornworts accumulated a list of questions that beauty of bryophytes. and some liverworts lack even a leaf- I might expect from people, who

VOLUME 31:3, JULY 2003 FREMONTIA 5 surprisingly want to continue con- chard is sparsely covered compared luted area.” My facetious answer elic- versing even after learning of my with such montane tropical its questions about bryophytes (and strange profession. For example, I rainforest trees. There, the trunk, lichens) and air pollution. Without am often asked, “Will the dense branches, and twigs may be cov- a root system, all parts of a bryo- mosses on my apple tree kill it?” And I ered to a depth of even a foot or phyte plant directly absorb their reply, “No, your healthy apple tree more, and they may even have dense water from raindrops and dew-fall. will not be hurt by its moss cover.” bryophytes on their evergreen Unfortunately the water that they This is because mosses lack a root leaves. In such conditions the absorb may contain dissolved sub- system, so they simply adhere to epiphyllous (growing on leaves) bryo- stances, including harmful or po- the bark of a tree but do not pen- phytes may damage coffee or cocoa tentially lethal materials from air etrate it. by interfering with photosynthesis. pollution. Vascular plants are bet- Only in rare cases will bryo- Having dispelled his worries ter protected from such direct air phytes have adverse effects. In the about apple trees, my conversant pollution damage. They take in so-called “moss forests” of mon- may next ask about the mosses on nearly all their water via the root tane tropical rainforests, bryophytes his roof. “Yes, they may damage roofs system, and those roots have a tis- may become so heavy as to pull by interfering with drainage.” “How sue (the endodermis) which excludes down a branch or even a tree dur- do I combat the moss cover on my roof?” harmful materials. ing heavy rains. Even an apple or- “Move your house to a more air-pol- “I have heard that lichens are used in mapping air pollution.” Histori- Clockwise from top: Lophocolea cuspidata. Most leafy liverworts have lobed or divided cally, bryophytes and lichens have leaves, whereas mosses never do. • Jungermannia sp., a leafy liverwort, with capsules. both been used for this purpose. Capsules in the leafy liverworts are relatively fragile and short-lived, and are normally Individual species of bryophytes and dark and spherical. • The leafy liverwort Gyrothyra sp. Leafy liverworts appear to have leaves in two ranks, and unlike mosses, the leaves never have a costa. lichens differ in response to air pollution. Some can be found in highly polluted areas, and others are killed by even a small amount of air con- tamination. This sensitivity to pollution is often a species-specific response to levels of mineral ions such as chromium, nickel, copper, sulfur, etc. Such species-oriented studies are more advanced in lichenology, and many cities, especially in Europe, produce pollution maps based upon the presence or absence of certain lichen species. Compared with lichenological studies, bryologists mapping air pollution are less concerned with species distribution around pollution sources. Instead they focus on mineral analyses of bryophytes in various air pollution regimes. Because mosses absorb all the minerals contained in incoming water, scientists in Europe have been hanging cloth- enclosed bags of mosses from free- way overpasses. Harvest of these bags at specific intervals is followed by mineral analyses. Interestingly, these studies show recent improve- ment in air quality in most cities in northern Europe. “Can I clean the mosses from my roof after they are killed by the drought

6 FREMONTIA VOLUME 31:3, JULY 2003 of summer?” Most mosses are not killed by drying. Without a root system, mosses must survive drying rather than merely resisting it. Flowering plants may use the rain- water of last month and even last year by continuously tapping the underground soil reservoir pen- etrated by their roots. Bryophytes, on the other hand, can only use the water of yesterday and today. Flow- ering plants resist the loss of water by covering leaves and stems with a waxy covering, the cuticle. While bryophytes may have a thin cuticle, it does not protect them from drying. In fact, protection from drying would run counter to their need for being able to absorb surface water. Because bryophytes must have plant parts capable of rapid water absorption, those same plant parts lose water with equal rapidity. Species of bryophytes differ in rapidity of water loss and water uptake, and these differences bear close relationship to the substrate on which they grow. Some bryophytes get virtually all of their water from precipitation (rain, dew, and mist). This is termed an ecthydric strategy, and the extreme of such a strategy is shown by bryophytes growing on the extreme branchlets of trees. In contrast, bryophytes of wet soil or logs exhibit a more endhydric strategy in that they typically take their water from the substrate surface and conduct that water throughout the plant in such capillary spaces as those formed by the wick of filamentous hairlike processes (rhizoids) on the stem surfaces. These two strategies are merely polar distinctions; in-between the extremes are innumer- able variations of water uptake depending on the substratum. Easterners have often asked me, “Why, in California, are there so many epiphytic mosses (growing on the trunks of trees)?” These epiphytic mosses are, of course, ecthydric, and such Conocephalum conicum (top) with sporophytes. • The thallus of Conocephalum conicum (bottom left) is several cell layers thick, with air chambers and air pores, as indicated mosses are completely hydrated by by the compartmentalization apparent in this image. • Ricciocarpus natans (bottom only a few millimeters of rain or right), a thallose liverwort. The photosynthetic tissue of thallose liverworts is thickened, even by water-droplet-laden fog whereas leafy liverwort leaves are usually one cell thick.

VOLUME 31:3, JULY 2003 FREMONTIA 7 eas of winter maximum rainfall (the forests, the south side of a tree may Mediterranean climate regions of even be limited by temperature ef- the west). In the East, the great fects. Bryophytes are largely cold- thunderstorms of summer are fol- weather plants with little or no pho- lowed by a hot drying period which tosynthetic effectiveness above soon leaves the mosses no better off about 80ºF. than before the rain. Winters in “Why did the moss on the rock in California have frequent periods of my front yard die when I sprayed it small and sustained drizzle or mist, with water last summer?” Mosses can and during those winters, the heavy be exposed even to the tempera- mass of epiphytic bryophytes shows tures of a Death Valley summer if most of its growth. they are dry and dormant. When In vascular plant ecology, we wet and warm, mosses will usually speak of hydric, mesic, and xeric plants use more food than they manufac- and plant communities. Such terms ture, resulting in starvation. do not work for bryophytes. It is “I was skiing the other day and better to speak of habitats on the noticed some thawing patches of snow. basis of 1) frequency of hydration/ Mosses and liverworts were protrud- dehydration, 2) duration of hydra- ing from that snow and apparently tion, 3) duration of dehydration, and growing. How can that be?” Most 4) intensity of dehydration. The flowering plants cannot realize a eastern United States has occasional net gain from photosynthesis at short periods of bryophyte hydra- temperatures much lower than tion after the rainstorms of the sum- 50ºF. It is true that some flowering mer, but these are quickly followed plants can grow through snow, but by long periods of dehydration be- that growth involves the use of food cause of the temperature of those manufactured and stored the pre- summer months. Winters in most vious year. In contrast, bryophytes of the East are characterized by fro- can often realize a net gain of pho- zen conditions without the possi- tosynthesis at temperatures very bility of fluid water necessary for near freezing. growth. In contrast, winters out- “Why are you crawling on the cold side the high mountains of Califor- winter ground?” My usual answer, nia allow for the continuous hydra- “I am testing the soil,” is the answer tion of the bryophytes, due to the of a coward fearful of the recruiters frequent mists and drizzles and low for the local mental institution. I evaporation of the cool winter am not really crazy. Many mosses months. and liverworts appear in late fall “Why do mosses grow on the north and early winter, and they dry to side of trees?” This is not necessarily invisibility in the summer. Such Mature hornwort, Anthoceros sp. (top). In this genus the spores are black, true! Mosses on the north side of bryophytes often grow and com- as indicated by the dark tips of the trees do not receive direct sunlight plete the lifecycle from spore to sporophytes. • Anthoceros sp. (bottom), with its heating and associated dry- spore in a period of 6–10 weeks. displaying both developing columnar ing effects. They may therefore re- They begin growth with the first sporophytes and gametophytes. The main hydrated and photosyntheti- moistening of the soil surface in gametophytes of hornworts can be distinguished from those of thallose cally active for a longer period of the autumn, and they must end liverworts by their blue-green, greasy time. Many trees have abundant growth when the grasses begin to appearance. bryophytes, not only on their north- shade them in the spring. Califor- ern base but also on all the rest of nia is a place of great diversity of banks. Most of the eastern United the trunk, limbs and even twigs. these ephemerals because many of States contains fewer epiphytic The total mass of bryophytes on a its regions have winters with un- mosses despite generally higher to- tree will be reflective of the dura- frozen soil surface left unshaded by tal rainfall. We are contrasting here tion of effective photosynthesis as vascular plants for several months the areas of summer maximum rain- opposed to the duration of drought- at a time. Northward, there is re- fall (eastern United States) with ar- induced dormancy. In more open duced duration of such unshaded

8 FREMONTIA VOLUME 31:3, JULY 2003 Variation in moss gametophytes (clockwise from top left): Hookeria lucens, recognized by its transparent overlapping leaves. In California, this moss is the species most likely to be mistaken for a leafy liverwort. • Buckiella undulata, a large pleurocarpous moss in California primarily found in the redwood forest zone. Photograph by J. Shevock. • Orthotrichum papillosum (with a few sporophytes). Orthotrichum is one of the larger moss genera in the state. • Homalothecium sp. This large pleurocarpous moss forms mats on rocks and tree trunks. and moist soil surfaces and with Conifers typically have the branches the water and minerals that flow that reduction we see reduced di- angled down from the point of in- down the trunks of trees. They only versity of ephemeral bryophytes. sertion on the trunk. Such a pattern slowly meter out that material to “I was in the redwood forests of allows the snow to slide off the the soil where it may be taken up northwestern California. The broad- branch tips. Without leaves during again by the roots of the trees. The leaved trees were drenched with mosses the winter, most of our broad-leaved loss of the moss mat from the trunks but the conifers were almost bare. trees can have branches upwardly of broad-leaved trees means that Why?” Conifers and angiospermous directed from their insertion on the the mineral nutrients enter the soil trees differ in many respects in the trunk. Such upwardly-directed too rapidly, and will therefore be habitats provided for bryophytes. branches allow water with contained lost into the water table and thence Conifer bark is typically more acid mineral nutrients to drain down to into the streams. Stripping mosses than angiosperm bark; it is usually the trunk and from there to the soil from tree trunks seriously depletes somewhat harder; and it continues at the tree base. The bryophyte the fertility of the forest soil. to be shaded during the high bryo- growing on a broad-leaved tree thus “If all that is true, why isn’t the phyte growth times of winter. gets more water and mineral nutri- soil of the coniferous forest rapidly de- Perhaps more important is the ents than the one on a conifer. Be- pleted of its mineral nutrients?” Co- greater amount of water and min- cause it gets more water, it grows nifers use and lose mineral nutri- eral nutrients provided to the epi- more, and can therefore intercept ents at a much reduced rate com- phytes on broad-leaved trees. Co- even more water and minerals. pared to broad-leaved trees. The nifer-branching architecture can be “Why are you so opposed to people low mineral content of water looked upon as an adaptation to the earning their living by stripping mosses leached from the canopy of a coni- breakage effects of snow load due from trunks of trees and selling them fer is spread around the forest floor to leaf retention during the winter. for floral designs?” Mosses intercept from the descending branch tips of

VOLUME 31:3, JULY 2003 FREMONTIA 9 10 FREMONTIA VOLUME 31:3, JULY 2003 from a tree and to tie it on to the tree with fish line. It is important that one take a fairly large patch because mosses operate on what I call a “Catch-22.” They can survive if there are enough of them to intercept and hold sufficient water during a dry time. In nature, one wonders how they get from being small and easily dried to becoming large and less easily dried. Another approach to planting bryophytes, and one that works especially well on rocks, is the use of plant fragments. Any living cell of almost any bryophyte has the potential to grow into a mature plant. A bryophyte can grow from frag- Variation in moss sporophytes: Opposite page, clockwise from top left: Buxbaumia viridis. Species in this genus are called “bug on a stick”! Buxbaumia is unique in that ments processed in a blender and you only see the plant when sporophytes are present because the gametophytes are so sprayed onto a suitable surface. But- inconspicuous. • Trachybryum megaptilum. This moss and other pleurocarpous mosses termilk is often included in the have capsules attached on lateral stems, as opposed to acrocarpous mosses, with capsules blender brew to provide a suitably attached at the tips of the stems. • Scouleria aquatica. At maturity, the capsules of this acid glue for these fragments. moss turn chestnut brown, and are shaped like an inner-tube. It is found seasonally submerged in rivers and fast flowing streams; when dessicated, the plants turn jet black. “Can I simply take the spores from • Polytrichum juniperinum, called the haircap moss for the netted coverings (calyptra) a moss capsule and tap them out onto a over the capsules. • Philonotis fontana is named the green apple moss, for its spherical suitable surface?” Probably not. Most capsules. Primarily found in springs and seeps. Photograph by J. Shevock. • Neckera bryophytes reproduce primarily douglasii. In California, this epiphyte is found in the coastal rainforest. Note the pendulous from plant fragments—whether sporophytes and ring of appendages (peristome teeth) at the mouth of the capsules. This page, left to right: Sphagnum sp. Members of this family (Sphagnaceae) are unique from the dirty toenails of squirrels among mosses: sporophytes lack peristome teeth and the leaves contain water-holding or the dried materials adhering to a cells and lack chlorophyll. • Heterocladium macounii. Note the peristome teeth. duck’s legs. Sporic reproduction seems to primarily be an adaptation that conifer. The high mineral con- fornia. I found maximum epiphytic to very occasional long-distance dis- tent of water leached from the bryophyte growth on years when persal, something that is probably canopy of a broad-leaved tree is there was little rainfall in the not very common. Experimental instead held in the moss mat on the normally wet months of October work in The Netherlands has shown tree trunk. In California those nu- through January. Minimum epi- that most bryophyte spores died trients are leached out of that moss phytic bryophyte growth occurred when exposed on the wings of air- mat, especially once the rains re- in times of extremely high late fall lines for even a single transatlantic sume in the late fall and winter. It and early winter precipitation. This flight. might also be noted here that late counter-intuitive observation seems As you might now more fully fall and winter, the time of maxi- to be related to excessive leaching appreciate, bryophytes are a fasci- mum leaching, coincides with the of the epiphytic moss mat with the nating group fully justifying my life- maximum growth of tree roots. consequence of reduced mineral long obsession. There is enough This results in more rapid uptake nutrients for growth. yet to be learned about these plants both through the root hairs and by “How can I grow bryophytes in my to justify an obsession of a similar the fungi that cloak the roots (myc- yard?” Japanese gardeners are fa- length, or a passion equaling mine, orrhizae). mous for their moss gardens, but for a few tens of thousands of ap- One of the most frequent com- such gardens are so labor intensive propriately curious amateur or pro- ments that I receive on a day of as to be daunting to a westerner fessional bryologists. heavy rain is, “A nice day for mosses [see Book Reviews, page 41]. More isn’t it?” My usual reply of “Maybe appropriate is the growing of mosses Dan Norris, University Herbarium, 1001 not!” is motivated by studies I con- on trees or rocks in your yard. Es- Valley Life Sciences Bldg., No. 2465, Uni- ducted in an area of temperate pecially with trees, it is often pos- versity of California, Berkeley, CA 94720- rainforests in coastal northern Cali- sible to take a large clump of moss 2465. [email protected]

VOLUME 31:3, JULY 2003 FREMONTIA 11 Sphagnum girgensohnii. Most people have heard of peat moss, which is the common name for members of the genus Sphagnum. Although peat moss covers large expanses of the northern hemisphere, and is ecologically one of the most important groups of mosses in the world, it is relatively uncommon and relictual in California. MOSS GEOGRAPHY AND FLORISTICS IN CALIFORNIA by James R. Shevock

osses are the largest of the Mediterranean climate. Only five The best answer seems to be that Mthree lineages of bryo- regions in the world have such a mosses are just the wrong size and phytes. With nearly 1,200 climatic pattern of cool wet winters have little or no recognized eco- species of mosses recorded for followed by long hot and dry sum- nomic value when compared to the North America, California has mers. The Mediterranean climate flowering plants. Our smallest moss nearly half of them. So what ac- is the defining factor in explaining in California is probably Ephemerum counts for this great diversity of the species rich moss flora in Cali- serratum that forms a green fila- mosses within the state? California fornia as well. mentous growth on soil after the is well known throughout the world Then why do plant enthusiasts winter rains. The capsules of Ephe- for its vascular flora, a flora influ- and professional botanists generally merum are around 0.25 millimeter enced and evolved in relation to a know so little about the mosses? in diameter. The other size extreme

12 FREMONTIA VOLUME 31:3, JULY 2003 is found in the genus Fontinalis, the water mosses, with plants up to one meter in length. Mosses are perceived to be hard to identify, have no showy features like flowers, and are just overlooked by many of us. But upon closer observation of our environment, mosses are actually all around us if we take a moment and focus our attention at a different scale. Viewed through a hand lens or microscope, mosses are simply stunning [For an introductory article to this group, see Fremontia 26(2): 3–8, 1998, sum- Ephemerum serratum (above left) is perhaps the smallest moss in the state of California, with capsules smaller than a pinhead, and is seen only during the winter months in marized on page 39 of this issue]. western North America. It is found on soil. Photograph by J. Game. • Fontinalis gigan- Mosses are no more difficult to learn tea (above right), commonly called the water moss. Plants are generally found submerged and identify than vascular plants once in streams and rivers, and can reach one meter in length. some basic new terminology and microscope techniques are learned. Mosses can grow on surfaces content to grow on rock surfaces, The major difference between that are generally not available to such as members of the genus mosses and vascular plants is that vascular plants. Mosses are also ex- Grimmia with over 30 of the 75 identifying most mosses to the spe- tremely important as pioneering species world-wide residing in Cali- cies level generally will require species as they can cover bare ex- fornia. Grimmia species can be greater magnification than can be posed soil caused by fire, landslides, found in every county of the state. obtained with a hand lens alone. or other perturbations. Even in ur- Other mosses prefer tree bark, and Another reason why the mosses ap- ban centers, mosses can grow on as a general rule, favor hardwood pear to be overlooked seems to be the sides of buildings, on concrete trees over conifers. Bark character- that botanists have placed an inor- retaining walls, and even asphalt istics like chemical composition, tex- dinate amount of value on being walkways! ture, and water-holding capacity are vascular at the expense of the rest of Mosses use a different strategy important in moss colonization of the plant kingdom. The result of to regulate water loss than do vas- this particular habitat. Mosses oc- this is that all of the plants and cular plants. Since they lack roots, curring as epiphytes on tree bark plant-like organisms without vas- they are not restricted to growing and branches do not adversely af- cular tissue are often simply lumped on soil. Many mosses are quite fect the trees. In fact, mosses have in botany textbooks as non-vascular plants, or cryptogams. Dendroalsia abietana (when dry, below left), a species endemic to western North America, We now recognize and acknowl- is a large moss, and especially common in the foothill woodland. When dry, the stems edge that mosses actually play a key curl, similar to a fern frond or the head of a violin. • Dendroalsia abietana (when moist, below right) with capsules arising from the underside of the branches. Photographs by role in ecosystem function, espe- J. Shevock. cially how they regulate the release of water and assist in protecting soil from erosion. Mosses can also tell us much about climate change. This is why the California Native Plant Society (CNPS) and botanists affili- ated with land management agencies are looking at the mosses in a new conservation light. To learn more about mosses from a non-technical approach, I highly recommend the recently published book Gath- ering Moss, a Natural and Cultural History of the Mosses by Robin Kimmerer. [See review on page 41.]

VOLUME 31:3, JULY 2003 FREMONTIA 13 blackburnpress.com) and will be of considerable interest to those want- ing to learn more about this fascinating group of plants. In addition, another great book to begin a study of bryophytes is Structural Diversity of Bryophytes by the late Howard Crum. It can be ordered from the herbarium at the University of Michigan, Ann Arbor. During the past 30 years or so the indefatigable bryophyte collec- tor Dan Norris has nearly doubled the number of mosses recorded from the state since Koch’s list. Dan’s collections, numbering over Ptychomitrium gardneri, an acrocarpous moss, found on rock. This is one of the few 106,000, come from all corners of mosses with several sporophytes arising from a single sexual bud (perichaetium). the globe and are permanently Photograph by J. Shevock. housed at the University of Cali- fornia, Berkeley. Probably many exploited just about every ecosys- of mosses. The botanists Bolander, CNPS readers are not aware that tem in California from sea level to Brewer, Bigelow, and Coville have prominent botanists such as Alice the summit of Mt. Whitney, from all contributed to this effort. In fact, Eastwood, John Thomas Howell, areas receiving barely an inch of Bolander was the principal collec- Willis Linn Jepson, Philip Munz, water a year in parts of Death Val- tor of mosses during the 1860s, and ley to the temperate rain forests many mosses new to science were Meesia triquetra (below top), a relatively dominated by coast redwoods. subsequently named for him to rare moss in California, found in Sierran Of course the lack of both iden- commemorate his efforts. In the fens. Photograph by J. Shevock. • Close- up of Meesia triquetra (below bottom), tification field guides or an illus- 1900s, other botanists collected showing the three-ranked leaves, as the trated moss flora for California has mosses in California, too. Leo Koch specific epithet implies. Photograph by not made it easy for plant conserva- and Fay MacFadden were the two J. Game. tionists, enthusiasts, or professional prominent bryologists who added botanists to learn about the mosses. many moss collections to herbaria However, plans are underway to in the 1950s. MacFadden was on provide such publications for our staff at Fullerton College, while state. Currently one has to use a Koch taught at Bakersfield College, variety of regional moss floras in Tulane University, and then at Illi- the United States to aid in moss nois. In 1950 Koch published his identifications. Lawton (1971) and list of California mosses in Leaflets Flowers (1973) are the two most of Western Botany, just after com- important moss floras available for pleting his PhD on the distribution California botanists. Dan Norris and of California mosses while at the this author have a catalogue of Cali- University of Michigan. fornia mosses and a key to species Other nationally known bry- ready for publication in Madroño that ologists during this era came out of is due to be published in early 2004. Stanford. William Campbell Steere spent much of his professional career at the New York Botanical HISTORY OF MOSS Garden, and Wilfred Schofield COLLECTING IN trained many bryologists while CALIFORNIA teaching at the University of Brit- ish Columbia, Vancouver. Wilf’s Many of the great botanical ex- Introduction to Bryology textbook has plorers of the 1800s in California recently been reprinted in a paper- were also involved in the collection back edition (available at www.

14 FREMONTIA VOLUME 31:3, JULY 2003 Peter Raven, Ledyard Stebbins, and Louis C. Wheeler collected mosses among their flowering plant work. However, their moss collections were generally few in number. Be- sides these prominent botanists, several moss floras at either county level or other administrative units were developed as part of master’s theses at the state universities at Humboldt, Pomona, and San Fran- cisco during the 1970s and early 1980s. Amateurs and botanists working within the land management agencies are primarily responsible for the current interest in the study of California bryophytes. Rarity and conservation of bryophytes are now being included as components of large landscape management strategies to provide for biodiversity at various scales in both the Pacific Northwest and in the Sierra Ne- vada. The Northwest Forest Plan, a major Environmental Impact Statement planning document established in the 1990s within the species range of the northern spot- ted owl, specifically requires sur- veys for cryptogams thought to be rare and restricted to old growth forests. These survey requirements facilitated a need for botanists to expand their scope of knowledge to include the identification of and survey protocols for bryophytes as well as other cryptogams.

Top to bottom: Andreaea rupestris. Members of this genus are commonly called the granite mosses, and are restricted to acidic rock types. The capsules on Andreaea are unique: note the four splits on the burgundy-colored capsule. In the field, the whole plants are usually about the size of a human thumbnail. • Aulacomium androgynum is a common species on logs and rotting wood. Note the spherical gemmae which are asexual reproductive structures, that are, simply speaking, a whorl of modified moss leaves. • Schistidium rivulare always occurs on rock, usually on seasonally wet areas such as boulders along a stream, or rocks of intermittent streamlets. The peristome teeth of the capsules are bright red to orange.

VOLUME 31:3, JULY 2003 FREMONTIA 15 MOSS FLORISTICS

While California lacks an illustrated moss flora, moss floras and checklists have been produced for the state going back to the 1860s. Leo Lesquereux presented the first paper on California mosses to the American Philosophical Society in 1863, and in 1868 the California Academy of Sciences published his Mosses From the Pacific Coast of North America. Mosses documented from California were prominently referenced in that work. Again, it was the moss specimens acquired by Bolander that contributed most to this undertaking. In 1880, the second volume of the Botany of California edited by Sereno Watson was published, and it now included the mosses. This was the first flora to address all of the mosses recorded or presumed to occur in California at that time. This moss flora documented 181 mosses for California. By the time of Koch (1950) the number of mosses documented for California had nearly doubled to 317. A new catalogue of California mosses referenced earlier by Dan Norris and this author will nearly double the number of mosses documented for California since the Koch publication. While mosses have been recorded for each of the 58 counties in California, some portions of the state have been more systematically sampled. In part, this reflects the location of educational institutions where bryology was taught in the past. The coast redwood forest region and adjacent coastal environments from Santa Cruz County north through Del Norte County have been fairly well sampled when Clockwise from top left: Hylocomium splendens, called the stair-step moss. Although compared to other regions of the abundant as a forest floor covering in North America, it is rare in California, restricted state. Moss catalogues, checklists, to the California-Oregon border. Photograph by J. Shevock. • Buxbaumia piperi, a or floras have either been devel- species seen only when sporophytes are present. Buxbaumia is generally found on logs, oped or are in preparation for Del and occurs in the northwest corner of California. Note the extension of the peristome Norte, Lake, Marin, San Francisco, teeth. Photograph by J. Game. • Racomitrium varium, commonly found on coastal rocks. The whitish tips (hyaline awns) of the leaves give this moss a frosted appearance and Santa Cruz counties. The mas- when dry. Photograph by J. Game. • Sphagnum capillifolium. This Sphagnum species, ters theses produced at Cal Poly although rare in California, is very distinctive due its reddish-tinged hue. Pomona in the 1970s focused on

16 FREMONTIA VOLUME 31:3, JULY 2003 other types of landscape units such In all of California’s deserts rain- while other species prefer a higher as mountain ranges or wilderness fall is considerably reduced, and pH like carbonate rock types in- areas within southern California. desiccation tolerance of mosses be- cluding marbles and limestones. But with the demise of many comes a more important factor, Perhaps the mosses most re- field-oriented university biology along with growth form, in under- stricted by substrate specificity are programs in the 1990s, all of the standing species distribution pat- the “copper mosses.” Both Mielich- campuses that once taught bryol- terns. Mosses in the families hoferia elongata and Schizymenium ogy in California have ceased to Pottiaceae and Grimmiaceae, with shevockii are restricted to rock types offer such training and education. dense cushion growth forms with high concentrations of met- It is through the efforts of Dan (acrocarpous mosses), are dominant als like copper and are primarily Norris as well as the Jepson Work- in arid environments, whereas located within the foothill wood- shop Series [see page 40] that cur- mosses that have a matted, pros- land areas. Scopelophila cataractae in rent taxonomic training in bryol- trate and highly branched growth California has only been collected ogy is being provided, much of this form (pleurocarpous mosses) are once in the Copperopolis area in training directed to expanding the indeed uncommon in the deserts. Calaveras County. Pseudoleskeella skills of amateur and professional Some of the desert mosses oc- serpentinensis, as the species name botanists, especially those employed curring in California also occur in indicates, is restricted to serpentine with land management institutions. other desert regions around the areas in northern California. world. In many cases, it is the same Other mosses like to occur along species, or a sister species occupy- rocky intermittent streamlets and MOSS GEOGRAPHY ing the same basic ecological niche. rock slabs with little competition. As one would expect, there are Mosses preferring this type of habi- Much has been written about fewer mosses in the dry interior of tat include Didymodon norrisii and the “California Floristic Province,” the state than in the coastal coun- Schistidium spp. Some mosses are an area not confined to the state ties. [See articles in this issue by nearly exclusively aquatic such as boundary but excluding the Great Lloyd Stark and Daniel Norris for members of the genera Fontinalis, Basin and Desert regions. The Cali- more details.] Hygrohypnum, and Scouleria, exposed fornia Floristic Province extends Climate and precipitation are only during low flows of unpolluted northward into Southern Oregon the primary factors influencing spe- rivers and streams. Although there near the Medford and Grants Pass cies distributions. For mosses, the are no marine mosses, some species region, and southward to the north- recurring interval pattern between like Schistidium maritimum can tol- ern portion of Baja California cell saturation and desiccation sets erate salt spray. Schistidium mari- Norte, Mexico. As already noted, the ecological tolerances for many timum occurs from Mendocino winter precipitation is a key feature moss species. Once cells are satu- County northward. Only a handful of a Mediterranean climate, and the rated, mosses have no ability to store of mosses can adapt to alkaline soils rainfall increases northward and additional water. Some mosses need that contain a high accumulation of westward across the California Flo- to be wet or moist most of the year, salts. ristic Province. such as species in springs and mead- Within the California Floristic Beyond the California Floristic ows or along rivers and creeks where Province, the distribution pattern Province, the eastern portion of they are seasonally submerged dur- of many mosses follows a general California is influenced by the Great ing peak flows. pattern that is observed in the flow- Basin Floristic Province dominated Secondary factors influencing ering plants. Common mosses in by sagebrush and pinyon pine-juni- species distributions include sub- the coastal northwest portion of per woodlands that are well repre- strate availability and elevation. the state become increasingly un- sented in Modoc and Mono coun- Mosses occur in California from sea common southward, making their ties. This last region includes the level to over 14,000 feet. Some spe- southern limit in the northern Si- Colorado and Mojave deserts, and cies are relatively common such erra. Many mosses of the coastal is dominated by creosote bush scrub as Bryum argenteum and Funaria counties such as Alsia californica, in portions of Inyo, Imperial, Riv- hygrometrica which have very wide Bryolawtonia vancouverensis, and erside, and San Bernardino coun- habitat parameters and can be lo- Triquetrella californica are not lo- ties. Therefore, it is fairly easy to cated throughout California. Oth- cated inland, just as many of the describe the pattern of moss distri- ers, however, are very specific to mosses located in the alpine zone of butions in the state by reviewing micro-environments. Some mosses the Sierra Nevada are not found the annual amount of precipitation prefer particular rock chemistry, elsewhere in the state. each area receives. such as acid rocks like granitics, Two regions of the state are

VOLUME 31:3, JULY 2003 FREMONTIA 17 after the first rains in the fall and can continue to grow until the last rains in May. The moss flora of California is also interesting when compared with the moss floras of adjacent states. Only about 50% of the mosses recorded for Arizona also occur in California. As with the flowering plants, water availability and climate play a key role in species distributions. The distribution of many moss species that occur in Arizona but not in California can be attributed to the availability of summer rain. In California, mosses are basically dormant in the summer, yet in much of Arizona, growth can occur in the summer due to that state’s mon- soon rainy season. Because of this climatic feature, the moss flora of Arizona has greater affinities with Mexico and the eastern United States than it has with neighboring California. For the most part, only the mosses of the Mojave and Colo- rado deserts have distributions which overlap both states. The state of Nevada is less known bryologically, but it has many mosses in the high mountain ranges that have greater affinities to the moss flora of the Rocky Mountains. In the Great Basin Floristic Prov- ince, winter growth is limited due to snow, and the region does receive some summer precipitation Schistostega pennata (top), commonly called the goblin moss. While not yet found in and this in part accounts for these California, this very small (less than 1 cm) but distinctive moss should be found in differences. Even Oregon, once be- areas with windthrow (exposed rootballs) and in openings of rock recesses, such as a yond the northernmost section of cave. The nearest location is in southwestern Oregon. • Pleurozium schreberi (bottom), commonly called red-stem, is one of the most common feather mosses (i.e., pleuro- the California Floristic Province, has carpous mosses) of North America. However it is yet to be found in northern Cali- a different moss flora. There are at fornia, where it should occur. least 70 species of mosses in Or- egon which have not been docu- noteworthy for the amount of moss the Sierra Nevada between the blue mented to occur in California. These biomass occurring in the ecosys- oak and black oak zone, especially species are more aligned to taxa tem. The first is the area domi- along the river canyons. Here many found in British Columbia and nated by the coast redwood forests. species of mosses carpet tree trunks Alaska, and most are generally Epiphytic masses of Isothecium and boulders, such as Antitrichia cali- pleurocarpous mosses. Several of the myosuroides and liverworts drape fornica, Dendroalsia abietina, and large and dominant mosses on the from the branches of trees, forming Homalothecium spp. In both cases, forest floor in much of western a tapestry of pastel greens while the these areas are basically snow-free North America such as Pleurozium forest floor is carpeted with mosses. although snows may linger for a schreberi have not been found in Cali- The second area where moss cover few days after major storms. Growth fornia. Hylocomium spendens, perhaps is abundant is along the foothills of can occur once plants are hydrated the most common pleurocarpous

18 FREMONTIA VOLUME 31:3, JULY 2003 moss species in North America, has main, and no Sphagnum have been matic era. Atractylocarpus flagellaceus only been recorded three times in documented from any of the moun- and Campylopodiella stenocarpa are California, and then only along the tain ranges of southern California. two such species that have recently Oregon border. Perhaps the most remarkable sta- been documented for California and Distributions of large moss gen- tion for Sphagnum in California is era are a fascinating study here in restricted to a small wet meadow Tetraphis geniculata (below top), is named California too. The genus Raco- surrounded by a pinyon pine wood- for the bent stalk (seta) of the sporophyte. mitrium and Sphagnum offer two land near Bodie, Mono County. Although reported for California, a specimen of this species has not been great examples on species distribu- Clearly this is a relictual occurrence, confirmed. Its sister species, T. pellucida, tion patterns. Mosses in the genus a holdover from the Pleistocene is fairly common in northwest California, Racomitrium, with nearly 100 spe- with no suitable habitat for miles in especially in the northwest’s redwood cies worldwide, generally inhabit all directions. The nearest occur- forest. This is the only moss in California rocky substrates. This genus is well rence of a Sphagnum species to this with four peristome teeth. • Tayloria sp. (below bottom) is a member of the dung represented in North America. site is nearly 4,000 feet higher up moss family (Splachnaceae). Currently With 18 species of Racomitrium for along the Sierran crest. this family has not been found in Cali- California, we see a rapid decrease Because mosses generally have fornia, even though Tayloria serrata in the number of species as one much larger geographical distribu- occurs as close as southern Oregon (near leaves the northwest corner of Cali- tions than vascular plants, the num- Crater Lake). fornia and the northern Sierra. By ber of mosses endemic to Califor- the time one arrives at the southern nia is therefore reduced. What we Sierra, over half of the Racomitrium have instead is a distribution pat- species have dropped out. This ge- tern of widely-spaced disjunct oc- nus is totally absent from mountain currences. Many mosses are re- ranges in the Mojave Desert and is corded from California from only rare in southern California. five to ten occurrences. In some Peat mosses in the genus Sphag- cases, this is merely a reflection of num offer a different distribution the number of herbarium specimens pattern in California. There are available for study and the collect- about 200 species of Sphagnum in ing habits of botanists, yet other the world and they are common in mosses are indeed very rare in Cali- higher latitudes, especially in the fornia. Some mosses in California Northern Hemisphere. The 23 spe- must now be viewed as relictual species of Sphagnum in California are cies—holdovers from a different cli- restricted to bogs or perennially wet meadows and fens and are distributed in two distinct patterns. Spe- cies of Sphagnum from the first group are coastal, found along la- goons or in areas with poor water drainage among the coast redwood belt in northeast California, with the greatest species diversity occurring at the pygmy forest region near Fort Bragg in Mendocino County. Sphagnum species from the other group are montane and are concentrated in the Sierra Nevada in meadows, fens, and lake shore habitats, from the lodgepole pine forests up to the alpine zone, generally in association with blueberry (Vacci- nium), labrador tea (Ledum), Kalmia, and sundew (Drosera). By the time one gets to Tulare County, only a handful of Sphagnum species re-

VOLUME 31:3, JULY 2003 FREMONTIA 19 calized extirpations by alteration of the “SO BE FREE” bryology for- habitat or loss of their specific mi- ays held annually during spring cro-environments. In many cases, break (see “Workshops” on page natural recolonization from such 40). SO BE FREE is an ongoing losses would be extremely unlikely series of West Coast forays orga- because available spore or plant nized by Brent Mishler at the Uni- fragments are so far away. In nearly versity of California, Berkeley. For any study area for mosses in Cali- more details and a review of the fornia, a suite of mosses will be past eight forays, visit the web at “rare” for that location. http://ucjeps.herb.berkeley.edu/bryolab/ While California has nearly 600 trips/sobefree.html. mosses documented to date, they It is hoped that many more are not evenly distributed among CNPS members will find the world the 58 counties in the state. Over of bryophytes rewarding and stimu- 300 mosses have been recorded for lating. There is a whole new world the southern Sierra, and even our of plant diversity just waiting for smallest county, San Francisco you to explore! County, has a moss flora of over 130 species. Probably the county with the richest moss flora will turn REFERENCES AND out to be Siskiyou County, but a FURTHER READING complete species list is not currently available. There are probably only Crum, H. 2001. Structural diversity of about 100 mosses for the entire bryophytes. University of Michigan Mojave Desert, yet this is the least Herbarium. Ann Arbor, MI. known area bryologically in Cali- Flowers, S. 1973. Mosses: Utah and the fornia, so our knowledge of this West. Brigham Young University. Provo, UT. portion of the state is bound to Kimmerer, R.W. 2003. Gathering moss, change once more bryophyte col- a natural and cultural history of the lections are made. mosses.Oregon State University Therefore, the current picture Press. Corvallis, OR. of the mosses in California is one of Lawton, E. 1971. (Reprint paperback). a highly species-rich flora with nu- Moss flora of the Pacific Northwest. merous species yet to be docu- Hattori Botanical Laboratory. mented. In addition, species of Nichinan, Miyazaki, Japan. mosses new to science are also be- Malcolm, B., and N. Malcolm. 2000. Mosses and other bryophytes: An illus- ing discovered in the state. trated glossary. Micro-Optics Press. CNPS members have a tradi- New Zealand. New species for California are being tion of adding distribution records Schofield, W.B. 1985. (Reprint 2001). discovered all the time. Pictured here is for our flora. For the first time, we Introduction to bryology. Blackburn Jim Shevock documenting the first have bryophytes listed in the CNPS Press. Caldwell, NJ. California occurrence of Helodoium blan- Inventory of Rare and Endangered _____. 2002. Field guide to liverwort dowii from Kings Canyon National Park. Plants of California. Much inventory genera of Pacific North America. Glo- Photograph by S. Haultain. • Helodium bal Forest Society & University of blandowii is a large pleurocarpous moss and fieldwork remains in Califor- nia. Current bryophyte workers like Washington Press. Seattle, WA. restricted in California to fen habitats. Shaw, J.A., and B. Goffinet. 2000. Photograph by J. Shevock. Colin Dillingham, Lawrence Bryophyte biology. Cambridge Uni- Janeway, Ken Kellman, Eve Laeger, versity Press. New York, NY. are distant disjuncts from central Ronald Robertson, and David Shevock, J.R. 1998. Bryophytes: Little Mexico. Toren are adding new species to forests among the trees. Fremontia The greatest threat to mosses in the state and filling in the distribu- 26(2):3–8. California is not species extinction, tion range for many species. There but rather, the loss of unique and are also opportunities for plant en- James R. Shevock, National Park Service, highly isolated occurrences. These thusiasts to learn more from either Cooperative Ecosystem Studies Unit, 337 isolated and small populations are attending a Jepson Herbarium Mulford Hall, UC, Berkeley, CA 94720. considerably more vulnerable to lo- Workshop course on bryophytes or [email protected]

20 FREMONTIA VOLUME 31:3, JULY 2003 Marchantia polymorpha is a common thallose liverwort of springs, seeps, creekbanks, and other wet areas. Pictured here with umbrella-like male branches bearing sperm-producing organs. THE ROLE OF THE AMATEUR IN BRYOLOGY: TALES OF AN AMATEUR BRYOLOGIST by Kenneth Kellman

am an amateur botanist. Like working on. I have been going on to over 400 species. In an effort to Imost members of the California like this for perhaps 25 years, and I make the list as complete as pos- Native Plant Society (CNPS), I have never thought that I would sible, I tried to reach every corner enjoy hiking in a local meadow or find anything that someone else did of the park. During one of those along a creek, identifying the flow- not know about . . . until I started hikes, I came upon a colony of leath- ers and shrubs that I find along the studying bryophytes. ery looking plants growing in the way. By trade I am a commercial air middle of a sandy creek bottom. It Each spring I run across old bo- conditioning mechanic, and my for- tanical friends, struggle to remem- mal training in botany has been Kindbergia oregana, one our most ber the names of the less common minimal, consisting of one class in common feather mosses. It is especially plants, and regularly find plants that plant systematics and one in den- prevalent on moist logs and soil banks on forest edges. Photograph by J. Shevock. I have never seen before. My pro- drology at the University of New cess for identification involves keys Hampshire in the early 1970s. I have and drawings in several references. no degrees nor have I ever been Often enough, I have to call my close to getting one. The rest of my mentor and describe the plant to knowledge has come from a persis- him. Usually he makes a sugges- tent curiosity, and the good will of tion, or asks a question that points more knowledgeable friends. me in the right direction, and I con- Sometime in 1992, I took on a tinue my search. After I feel com- project to compile a list of the plants fortable with an identification of a in a local county park where I was a plant, I make a note in my local docent. It was a small park of 300 flora, or in my Jepson Manual, or acres in Santa Cruz County, filled perhaps add the name to a list I am with microhabitats, and my list grew

VOLUME 31:3, JULY 2003 FREMONTIA 21 I thought I would only show I knew that I did not possess him this one colony, but inside of some special attraction to rare two hours, he had not only identi- mosses; I was just collecting every- fied my plant (Marchantia poly- thing that I found. I also learned morpha L.), but had also shown me that day that the distribution of ten other species of liverworts and mosses was poorly understood in hornworts! I had never heard of this most populous state in the na- leafy liverworts, and had no idea tion, and that I could help correct that they were abundant on the bark that. of local rocks and trees. I thought I So I continued collecting and had been looking very closely at identifying my collections, but Dan this park, yet I had been missing an Norris had moved to Washington entire suite of species and their habi- State and I lost his expert assis- tat. That was the day I was bitten by tance. As a neophyte, I did not feel the bryology bug. It was a dream confident in my keying decisions, come true for any naturalist . . . a and without Dan’s confirmations whole new world to explore. and corrections I floundered for a Bill very graciously taught me while. Then in the spring of 1998, how to collect and then helped me Jim Shevock wrote an article on identify my collections. He gave me bryophytes (Fremontia Volume keys, and allowed me to copy de- 26(2):1-8). I contacted him and scriptions from antique books that within six months or so, my “re- he owned. He has continued to help search” rose to a new level. me to this day. Jim wanted to cite my collec- Soon I bought my own micro- tions in a paper he was writing with scopes. While my work with vascu- Dan on the distribution of Califor- lar plants continued, I was now rec- nia mosses. I had been working on ognizing the more common species a list of Santa Cruz County bryo- of liverworts and hornworts on my phytes, but never thought it would Overall aspect of Leucolepis acanthoneura hikes. After working on these plants go beyond that. Jim encouraged me (top), a moss found along moist stream- for about nine months, I enrolled to start housing my collections in sides and shaded forest areas. • Individual in the Jepson Herbarium weekend an herbarium, and told me that my plant of Leucolepis acanthoneura (bottom) workshop on bryophytes. Two very work was significant enough to pub- showing “palm-tree-like” growth form. The bracts on the stem are white; leuco- important things happened as a re- lish in a scientific journal. Taking lepis means “white scale.” Photographs sult of that seminar. First, I was Jim’s suggestions to heart, I de- by J. Game. introduced to the study of mosses. cided to write a “Catalog of the Until that time I had purposely ig- Mosses of Santa Cruz County,” and had spectacular little umbrellas nored mosses while I concentrated to submit it for publication. springing up from the thallus, and on the liverworts. Second, I met Eventually, Dan returned from little fringed cups, filled with “eggs.” Dan Norris, who was teaching the Washington, and he, Jim, and I met I had a vague memory from my bryophyte workshop. every couple of months to review systematics class that this was a liv- Dan provided us with glossa- my collections and work. I combed erwort, but I had no idea what spe- ries, keys, and bibliographies to get through the collections in the three cies it was. I went home and started us started, and he encouraged us to Bay Area herbaria, noting all of the looking in my books, but none of collect. I was off and running. Three collections from Santa Cruz my references even mentioned liv- months later, I brought him a box County. This review of existing col- erworts. I called my mentor, and of perhaps 150 packets of mosses. lections pointed out exactly how asked him, but this time he was He patiently went through them little was known about bryophyte only able to give me someone’s and gave me tentative identifica- distribution here. name to call. Eventually I was re- tions. But in that box was one moss Of the approximately 190 spe- ferred to Bill Doyle at the Univer- that had never before been collected cies of mosses that I have docu- sity of California, Santa Cruz. I was south of Humboldt County. It was mented in Santa Cruz County, well delighted when he offered to meet on that day I realized that an ama- over half had never been collected me at the park and look at these teur bryologist could make a “con- from the county before. Fifteen of liverworts. tribution to science.” those have never even been col-

22 FREMONTIA VOLUME 31:3, JULY 2003 lected from the central coast of Cali- fornia. Seven of those 15 are still undescribed species, and local collections like mine add to the knowledge of range and morphological variation for each one. I still collect, and even after eight years, I am still finding new taxa to add to my list. My moss catalog has been published in Madroño (Volume 50:2), the journal of the California Botanical Society, and I am starting a formal catalog of the liverworts and hornworts of Santa Cruz County as well as starting collections in Monterey County for a bryophyte catalog there. Collecting bryophytes is like a grand treasure hunt, and it is a great excuse to get outside and enjoy nature. Publishing a local catalog forced me to collect throughout the county, and so I have been to beautiful creeks and forests that I had never visited before. Another plus is that bryophytes (mosses in particular) grow everywhere—on dry rocks, in the water, on soil, on cement, even in the city—and they can be collected any time of the year. They are very easy to collect and store. No pressing or mount- ing is allowed, much less required. Just put the plant into a packet, note the collection data, let it dry, and voila! This is nothing like the amount of work required for collecting a vascular plant. The packets are small enough that I can keep a hundred specimens in a bureau drawer. Since these plants desiccate in nature, a dry moss in a packet looks exactly like a dry moss on a rock, and because mosses preserve

Isothecium stoloniferum (top) is a common epiphytic moss in the temperate rainforests in northwest California. • Anti- trichia californica (middle), a widespread species. Note the closely associated, concave leaves which give a wormlike (julaceous) appearance to the leafy plant. • Pseudobraunia californica (bottom), another julaceous moss. Pseudobraunia forms large bronze-hued mats on rocks. Photograph by J. Shevock.

VOLUME 31:3, JULY 2003 FREMONTIA 23 Studying bryophytes is not only understood in the most populous exciting for the amateur, it is im- state in the US, but it is true, and it portant. Most academicians have will be true for some time to come. abandoned bryophyte floristics be- This provides the amateur natural- cause it does not attract funding. ist with an opportunity to make a Because of the discontinuous na- significant contribution to the ex- ture of bryophyte distribution (see isting body of botanical knowledge. Jim Shevock’s article on page 12), There are many opportunities bryologists aren’t even sure what for amateur bryologists to make a plants are rare in our state. It is difference in our understanding of difficult to accept that a group of these plants. There is, of course, a plants this large can be so poorly floristic project like mine, which

Plagiomnium insigne has some of the largest leaves of any Californian moss. It is found on shaded and moist soil and humus along springs, streams, and waterlogged areas.

Dicranum howelli (top) is widespread in northwestern California, and is primarily found on forest litter. • Atrichum selwynii (bottom), common on mineral soils, is an acrocarpous moss readily recognized by its undulate leaves with prominent costa. Photograph by J. Shevock. so beautifully, identification can be done the minute you get in from the field, or if that is inconvenient, in 25 years. All that is required is rehydration from a drop of warm water.

24 FREMONTIA VOLUME 31:3, JULY 2003 involves making a list of bryophytes and some of the basic bryophyte in a park or even an entire county. biology, people still love to see these Most of the counties in California tiny plants on a wildflower hike. represent giant holes in our knowl- How do you get started? I heart- edge, waiting for someone to make ily endorse attending the annual concentrated collections. Other bryophyte weekend workshop county studies have revealed spe- sponsored by the Jepson Her- cies new to California, and even barium. This hands-on class pro- species new to North America. vides a great foundation for your Without extensive collection it is studies, and includes collection and impossible to know what is rare and dissection techniques, recognition what is common. Creating a local of the various bryophyte groups, catalog is an excellent way to learn practice using keys, and an over- bryophyte identification. If a cata- view of bryophyte biology. (The log seems too ambitious a project, next Jepson bryophyte workshop just making collections is a valuable is February 21-22, 2004. For de- contribution. Local herbaria are al- tails see http://ucjeps.berkeley.edu/ ways hungry for properly identi- jepwkshp.html, or write to the Jepson fied, well-labeled specimens. Herbarium, University of Califor- For those with a camera and a nia, 1001 Valley Life Sciences close-up lens, there is always a need Building, #2465, Berkeley, CA, for quality photodocumentation. 94720-2465.) Photographing bryophytes is noto- Another source of good infor- riously difficult, and will become mation and contacts is the bryo- valuable as interest grows and field phyte listserve on the Internet. To guides come into demand. Likewise, join, send an e-mail, with the single technical artists could draw bryo- word “subscribe” in the body of phytes for articles describing new the message to [email protected]. species, or for a small local flora. The annual bryophyte foray “SO Cataloging herbarium collec- BE FREE” sponsored by Brent tions is an invaluable service that Mishler’s laboratory at UC Berke- can be provided especially by those ley is a great event for amateurs to who enjoy working with comput- meet other bryologists of all skill ers. Very few herbaria in our nation levels. Information on the event is are yet part of computer databases, posted at http://ucjeps.berkeley.edu/ so volunteers can make a huge dif- bryolab/trips/sobefree.html. ference here by entering data into As your interest and commit- Top photograph: Dan Norris (left) and computers. Once collections are ment grows, you will need access Jim Shevock (right) during a Jepson part of computer databases, this to microscopes and a good refer- workshop in the Sierra National Forest, wealth of information suddenly be- ence library. Used microscopes can near Minarets pack station. Photograph comes accessible to people through- be obtained from local microscope by S. Markos, courtesy of the Jepson out the world via the Internet. A technicians for a reasonable price, Herbarium, UC Berkeley. • Bottom photograph: Dan Norris (left) and Jim bryologist studying a particular and investing in a personal library Shevock (right) working in the University taxon could then query a database is well worth the expense. Cur- Herbarium bryophyte collection, UC and request loans from the her- rently, there is no “Jepson Manual” Berkeley. Photograph by B. Ertter. barium, saving countless hours for California bryophytes, so a pouring over specimens. number of floras from North to help you if they sense a commit- Public education also offers the America and beyond must be used ment to learn. amateur additional opportunities to for identification. I am the first to I encourage you to polish your advance the field. Slide shows and admit that this work involves a long hand lens, and then open your eyes hikes that focus on bryophytes are a learning curve, so finding yourself to the tiny world that you have been great way of introducing people to a mentor is almost imperative. But walking by every day of your life. these wonderful plants. Even if your the bryologists I have met have knowledge is limited to recogniz- proven to be a friendly and gener- Kenneth Kellman, 9870 Brookside Ave., Ben ing the major bryological groups ous lot, and will be more than happy Lomond, CA 95005. [email protected]

VOLUME 31:3, JULY 2003 FREMONTIA 25 MOSSES IN THE DESERT? by Lloyd R. Stark

hen I first came to the briefed me on what to look for based call very distinctly when the pas- WMojave Desert as a stu- upon the northeastern California sion to study mosses returned. A dent intern in botany, it “high desert” region. short trip into a desert canyon near was so hot and dry that I was amazed Many years passed before I took Lake Mead allowed me to survey that plants, least of all mosses, could up a permanent residence in the the area for mosses, fully expecting survive the heat and scarcity of rain- Mojave, and at this point I had dis- to find very little in the way of these fall. After all, mosses require a moist tanced myself from a study of plants. After all, generations of environment to grow and repro- mosses. Although I had spent some bryologists had routinely avoided duce, and the Mojave Desert is the time in the Chihuahuan Desert of the desert with the preconceived hottest and driest region on the con- southern New Mexico, when I came notion that it represented a vast void tinent. Nevertheless, my instructor out to the Mojave Desert of south- harboring few if any species able to in botany at Humboldt State Uni- ern Nevada and California, I hon- tolerate the heat, lack of water, wind, versity, Daniel Norris, had con- estly did not expect to encounter and general severity of weather. vinced me that mosses did indeed much in the way of mosses in the Botanical collectors of flower- inhabit the desert, and he had lower elevation desert terrain. I re- ing plants in the desert often re- strict themselves to a few weeks of Typical Mojave Desert habitat for Crossidium species. A wash runs through the center collecting in order to avoid the heat. of the photograph (covered with green vegetation), and Crossidium is found at the base After spending nearly 15 years in of the small rocks on the north-facing side of this wash. Photograph by L. Stark. the eastern temperate forests where mosses and liverworts are abundant, and then relocating near Las Ve- gas, I can comment on this prevailing viewpoint of mosses being generally absent in deserts: it is utterly false. To my surprise, this short hike revealed several species of mosses, at which point my interest in these little plants was rekindled. So struck was I with this discovery that my career path actually changed at that moment, and I was drawn back into the field of bryology. This was about nine years ago, and I have never looked back. The mosses of the Mojave Desert have offered an out- door laboratory heretofore un- tapped, and have yielded some of the most fascinating patterns of reproduction and survival among plants. Now, strictly from the stand- point of diversity (the number of species in a given area), it is true that there are fewer species in the Mojave Desert than surrounding areas; Jim Shevock is estimating about 125 species as of this writing. While this number represents a small proportion of the estimated 600 species of mosses in California,

26 FREMONTIA VOLUME 31:3, JULY 2003 the desert species are perhaps more THE WAITING GAME interesting because: 1) several globally rare endemics occur in deserts, How are these rootless plants even a new liverwort awaiting de- that have no water storage capacity scription that will be named after able to survive the rigors of the the Mojave Desert; and 2) the spe- desert, where rainfall is not only on cies inhabiting the desert exhibit the order of millimeters per year, some amazing survival strategies of but is more sporadic and unpre- desiccation tolerance and reproduc- dictable than any region on earth? tion found nowhere else in the Mosses have no roots, no water stor- world. age organs, few protective pigments Although qualifying definitions to diffuse the sun’s rays, and occupy of deserts vary widely in the litera- only the surface of the substrate on ture, the boundary between desert which they grow. Therefore, they and semidesert can be recognized dry out as rapidly as the soil dries as occurring below the 120–150 mm out, which can range from a few isohyet of annual precipitation. By days following a winter rain, to a this definition, only the western few hours following a summer rain. edge of North America’s deserts In the Mojave, most of the rainfall conforms to a true desert, namely is during the cooler winter months, the Mojave and western Sonoran so herein lies a major irony insofar deserts (inclusive of the Colorado as desert mosses are concerned: they Desert of Southern California). are shade-adapted plants that do Compared to other deserts of the best when temperatures are cool and world, North American deserts are moisture is readily available. not only small in area, but youthful Optimal growth conditions for Four desiccated plants of Pseudocrossidium in age. As judged by paleobotanical desert mosses surprised everyone. crinitum (top) from the Mojave Desert. evidence using packrat middens, They prefer it to be 60–70ºF un- Leaves of desert mosses often show a char- present floristic compositions ex- der very dim light (full shade). I acteristic spiral twisting of the leaves about tend only to circa 8,000 years ago. struggled for years trying to grow the stem. • A wetted patch of Pseudo- crossidium crinitum (bottom) from the The boundary of the Mojave Desert desert mosses in a growth chamber Mojave Desert. This hydrated condition can generally be said to follow the until an undergraduate student dis- is evident only a few days following a rain- distribution of creosote bush (Larrea covered the plants have to be storm. Photographs by L. Stark. tridentata) and also the Joshua tree shielded even from the direct light (Yucca brevifolia). of a grow light! From April through know just how long a desert moss November, mosses in the desert tol- can survive in a dried state. Desert The desert biological crust of the Mojave erate the heat and desiccation in a mosses are photosynthetic oppor- Desert. Black patches on the soil surface dormant state. During this period tunists: they capitalize on the win- are composed of mosses and lichens, with of blistering heat, the cells do not dow of opportunity, from about the dominant moss Syntrichia caninervis. Photograph by L. Stark. die, but remain physiologically alive December to March, when a suffi- yet totally inert (respiration is un- ciently hydrating winter rain oc- detectable!). curs. Within hours of a rain they Going nine months without are carrying out positive net photo- water is a trick that not even the synthetic gain—making sugars. most heat tolerant Mojave Desert Thus the life of a desert moss is flowering plant can carry out. How- really a waiting game, where the ever, this feat is relatively easy for a plants patiently await a winter storm desert moss, most species of which that may only occur every few years. are able to tolerate several years of Recently I followed a few popula- total desiccation and then spring to tions of the most desiccation-toler- life when wetted. After eight years ant moss in the Mojave Desert, a in dry storage in the dark (in an species in the genus Crossidium, for herbarium), one specimen, to our a period of one year in order to amazement, showed 100% viability determine the duration of the hy- among its stems, and we still do not drated versus the dormant dry state.

VOLUME 31:3, JULY 2003 FREMONTIA 27 habitats. About 80% of moss species are bisexual in true desert habitat, compared to an expectation of 40–50% of mosses at large. Bisexu- ality allows the male and female reproductive structures to be in close proximity, allowing self-fertilization to occur, and thus the plants reproduce sexually by producing large quantities of airborne spores. In mosses—as described by Daniel Norris in his article on page 5—sperm is transferred from male to female reproductive structures by droplets of rain. The sperm is carried to the egg in a rather hap- penstance manner, having no directed vectors of dispersal (unlike pollen, which can be transported effectively by insects and even wind). Therefore, for a dioecious species, in which the male and female sexes occur on separate plants, the male Typical moss life cycle, showing diploid stages (zygote and sporophyte) and haploid and female plants have to be imme- stages (spores, protonema, gametophyte, and gametes). Illustration by L. Vorobik. diately adjacent to one another (within a centimeter), which seldom During this year (2001), the area peratures following a summer storm occurs; thus the selective advantage around Las Vegas received slightly may be 90–110ºF, and the moss finds of bisexuality. more (120 mm) than its normal an- its metabolism accelerated and un- The second most noticeable cir- nual rainfall (104 mm). These plants balanced, outside of its optimum cumstance regarding reproduction were fully hydrated for a total of 31 range. Photosynthesis is difficult and in desert mosses is that in dioecious days in 2001, partially hydrated (in substantial damage can occur that is species the sex ratio is exceedingly the process of drying down) for a unrepairable. Many plants and all skewed in favor of females. Further total of 35 days, and entirely desic- reproductive structures can die in a investigation revealed that this ra- cated for the remaining 299 days. matter of hours. Fortunately, sum- tio represented the most biased sex During one stretch these plants tol- mer thunderstorms are sporadic in ratio of any plant, flowering or oth- erated 147 consecutive days of des- the Southwest US, occurring about erwise. How biased you might ask? iccation. Fully wetting these plants once per year. How about: on the order of one required about 4 mm of precipita- male for every 100 females, or even tion, as rain. the total absence of the male, i.e., The mosses, then, “wait” for a BIZARRE entirely female populations. There good winter rain, after which, due REPRODUCTION are even a handful of species in the to the lower temperatures of winter, IN THE DESERT American Southwest in which male they can remain hydrated for several plants are unknown to science. On days. The first day of hydration is I was fortunate enough to be the contrary, there are no male- spent repairing their tissues from the exposed to an unexplored frontier, only species in this region. damage of the wet/drying cycle of to be the first one to really take a Finding such skewed sex ratios events (primarily membrane dam- look at how desert mosses repro- in desert mosses was most unusual, age). From the second day onward, duce. It is an ongoing privilege that because the expected sex ratio is they can grow and reproduce. It is in has consumed many pleasurable 1:1. Sex is chromosomally deter- their patterns of reproduction that hours of my time. One thing that mined, with the male plant carry- many an interesting tale can be told. became very noticeable was that ing a single y-chromosome, and the If a summer thunderstorm hits, it is many desert mosses were bisexual, female plant carrying a single x- very bad news to the mosses. Tem- more than expected in true desert chromosome. Equal numbers of

28 FREMONTIA VOLUME 31:3, JULY 2003 Mosses of watercourses (clockwise from top left): Didymodon brachyphyllus is seen here in a watercourse. This species is a member of one of the more difficult genera of mosses. Here it intercepts water and sand brought down in this stream channel during occasional storms. Photograph by J. Shevock. • Funaria hygrometrica. This may be the moss most frequently used in physiological experiments. It is abundant throughout the world, but is especially abundant on the ash of old burn piles. • Crumia latifolia is seen here in another watercourse. It belongs to the same family as Didymodon. Crumia has thickened leaf margins that may resist tearing during floods. • Fissidens bryoides. Fissidens is a very large genus primarily found in tropical areas. The flattened leafy stems (as indicated by the leaf arrangement) may remind one of a leafy liverwort, but the presence of a midrib in the leaves defines it as a moss. This moss often grows in wet seeps. • Bryum capillare is one our most widespread mosses in deserts and in mesic areas outside of desert regions. It is a member of the largest genus of mosses in the world. In deserts, it is found in seeps that have little or no salt accumulation.

VOLUME 31:3, JULY 2003 FREMONTIA 29 Mosses of sunny rock outcrops: (this page, clockwise from top) Grimmia laevigata is like most species of the genus in having hyaline awns (transparent leaf tips). These may give a frosted appearance to the colony, and thus insulate the plants from the heat of the desert sun. Photograph by J. Shevock. • Grimmia pulvinata is one of our most widespread members of its genus. It is especially common on cement blocks or walls in urban environments. It is found not only in deserts, but also in humid areas of the coast. Photograph by J. Shevock. • Grimmia moxleyi is the most desert- adapted member of this genus. It is male and female spores are pro- dispersal unit—a seed or spore) is restricted to the very hot and dry rocks duced, and one expects to find males very rare among dioecious desert of California’s southeastern deserts. Photograph by J. Shevock. • Opposite and females in equivalent numbers mosses. One such species, Syntrichia page, top to bottom: closeup of Grimmia in the field. In all desert dioecious caninervis, dominates the soil in moxleyi. This moss has hyaline awns species examined, the pattern of fe- zones of blackbrush, Coleogyne restricted to the leaves which surround male-bias is easily observed. Deter- ramosissima. Acres upon acres of fe- the sporophytes. When dry, the living mining the reasons why males males occur, with but a male here material of G. moxleyi plant cells is protected by black, opaque cell walls should be so much rarer than fe- and there, if the male is present at that shield cell contents from the sun. males has occupied the time of sev- all. In fact, we have yet to find a Photograph by J. Shevock. • Coscinodon eral scientists over the last few years. population that is exclusively male, calyptratus is another member of the One of the puzzles is that sexual while exclusively female populations Grimmiaceae, separated from Grimmia reproduction, with all of its sup- are commonplace. by the large and basally-lobed calyptrae (sporophyte caps). This moss, in Cali- posed advantages to a species (i.e., Successful sexual reproduction fornia, is restricted to the desert or desert enhancing genetic variation, repair- is hindered by several factors, fringes. Photograph by J. Game. ing the DNA, and producing a nice among them: 1) male rarity, 2) low

30 FREMONTIA VOLUME 31:3, JULY 2003 ently resources are so limiting (es- WHY ARE MALES SO pecially water and secondly nitro- RARE? gen in deserts) that the maternal plant does not have sufficient re- Male rarity is the primary rea- sources to dedicate to her offspring. son why sexual reproduction is so So, instead of sacrificing her own infrequent in dioecious desert potential to survive, she jettisons mosses, but why are there so few her offspring in order to preserve males to start with? Some low-el- the possibility of future reproduc- evation desert mosses are entirely tion for herself. After all, if she female, with the male of the species should die while nurturing her off- only occurring at higher elevations spring, she will not be able to re- where conditions are moister. Al- produce in the future. However, if though this sex ratio may sound she aborts the offspring due to a like a great situation to be in for the shortage of resources that she can males, this is really not the case at provide, she lives to reproduce an- all. The vast majority of males never other day, perhaps more than once. father any progeny, and >99.9% of A “decision” is made early on in females are virgins for life, i.e., a order to conserve resources. As a sort of chaste society if you will. result, nearly all abortions occur in The female-biased sex ratio in the embryonic phase. One nice mosses is quite opposed to the of- thing about studying reproduction ten male-biased sex ratio in seed in mosses is that a person can exam- plants. ine a female stem under a dissecting How can this be? In seed plants, microscope and see what happened it is the male that is more common, sex expression, 3) the highest known in each of the last three to four more vigorous, and more tolerant abortion rate among mosses, and 4) years of reproduction—reproduc- of xeric conditions. The prevailing local weather variation that causes tive structures (sex organs and abor- hypothesis for male dominance reproductive cycles to skip years. tions) usually remain intact along among seed plants states that it boils Low sex expression is characteristic the stem. down to energy budgeting: since of dioecious species of mosses in Rainfall in the Mojave Desert is the female has to nurture an ener- general, and desert species in par- one of the most erratic and unpre- getically expensive fruit, she then ticular. By that is meant that, if one dictable events on the continent. has less energy and resources to fight collects a few stems of this Syntrichia, Weather forecasters are not to be infection, survive environmental hydrates the stems on a microscope relied upon. In the winter of early stresses, and to devote to growth. slide, examines them under a dis- 2002, rain was predicted in the Las The male, on the other hand, pro- secting microscope, and probes Vegas Valley on five occasions, and duces pollen, which is cheap com- around the leaves and at branch nary a drop fell. This is a typical pared to fruit. He can afford to de- junctions in search of sex organs, in scenario. Since the mosses depend vote more resources to surviving all likelihood the stem will be found upon external water from rain show- the rigors of the desert. to be a “nonexpresser” (of undeter- ers for their growth and reproduc- With the mosses, however, the mined sex). tion, and really have a “window of tables are turned, and the male de- On average, less than 10% of all opportunity” that extends from votes more energy to reproduction. stems have expressed sex (produced about December through March, The cost of a sperm packet far ex- sex organs) over their entire lifetime. should insufficient rains fall during ceeds the cost of a single egg. A The average age of an individual this period, the entire reproductive male will produce hundreds of stem is about 15 years in the Mojave, cycle will be skipped. This exact sperm while a female produces only although clones can be much older. thing happened across the Mojave a few eggs. Because fertilization re- Even on the rare occasion when sex Desert for three consecutive years quires both sexes to be side by side is achieved, the odds are that the in the late 1990s. A few years ago, in the presence of external water, offspring generation resulting from we had a stretch in the eastern fertilization is so rare that females sex (called the sporophyte, a spore- Mojave of over 200 consecutive days do not experience their full cost of bearing structure) will abort. In fact, without a drop of rain. The mosses reproduction, producing only un- about 60–70% of all fertilizations lived on, but no sexual reproduc- fertilized eggs and no offspring to abort. Why is this the case? Appar- tion was possible. nurture (clearly, sex in the rain is

VOLUME 31:3, JULY 2003 FREMONTIA 31 not what it is cracked up to be). storm, you should be able to locate eas of heavy grazing, even after the Meanwhile, males always experience healthy populations of mosses that grazing has stopped. their full cost of reproduction appear a beautiful green. The scent The presence of high mountain (sperm) regardless of the presence of creosote bush from the sur- ranges in deserts creates ecological of a female. rounding lower elevations should elevation gradients over which plant Females, then, have a surplus of be sufficiently intoxicating at this assemblages change drastically resources to devote to desiccation time, and that should be your cue. within short distances. The lower stress, survival, growth, and main- Look for places at the bases of boul- tree line is encountered as one en- tenance, whereas males are com- ders and shrubs that receive no di- ters pinyon-juniper woodland, and promised in these activities. As a rect sunlight, even at high noon. the “desert” is left behind. Above result, the male has fewer resources The soil here remains moist for this point, at least five well-defined to devote to survival, desiccation longer periods, and is conducive vegetation zones occur, including a tolerance, and growth. The female, for mosses to grow. Once you get mesic evergreen forest at the high- then, is expected to grow faster, ex- the idea of what to look for, you est elevations. Thus the southwest- hibit greater clonality, and to oc- may want to venture into lower ern deserts contain a variety of cupy more desiccated habitats than elevation desert canyons and try nondesert communities. Across is the male. True enough, one rarely your luck. these seed plant communities, the finds males of desert mosses in true A sharp contrast exists between moss communities change too, and desert habitats (lower elevations) in north-facing and south-facing slope you can find a completely different the Mojave Desert. aspects: bryophytes may be absent set of mosses at high elevation than entirely from surfaces along a just a few miles down the road at south-facing aspect that exceeds 20 lower elevations. The higher in el- WHERE TO FIND degrees, whereas rich moss com- evation you go in deserts, the greater DESERT MOSSES munities can occur on north-fac- the diversity and prominence of ing sloping soils or, more com- mosses. It takes “an eye to the ground” monly, rock outcrops. The reasons Mosses can also be found in to perceive these little plants, but for their absence from south-fac- spring/seepage areas depending on with a hand lens you can find mosses ing aspects is called the “carbon water availability, water chemistry, in nearly every corner of the desert. balance hypothesis”: mosses on and surrounding vegetation. Here, In the desert, mosses appear much south-facing slopes dry out quicker too, cattle and burros have impacted like the color of the sand and rocks than mosses on the other side of the bryophyte community. The they inhabit, even blackish brown. the canyon due to warmer tem- desert offers the botanist or botani- However, upon wetting, these dark peratures and greater light expo- cal enthusiast the opportunity to mosses turn brilliant shades of sure. Mosses must be wet to photo- view a variety of habitats all within green. At lower elevations, mosses synthesize, and need to be moist a few miles of one another. By driv- are restricted to the north-facing for at least a full day after each ing a road from a lowland desert sides of outcrops and shrubs, where rainstorm to make sufficient sugars region up into a mountain range, they enjoy the shaded shadows and for growth and reproduction. one encounters a range of plant remain hydrated for hours longer Therefore, those plants on the communities, from creosote bush than adjoining exposed sites. In south-facing side of the canyon can- to ponderosa pine. desert canyons, it is possible to come not reach a positive carbon balance As you might expect, the changes across carpets of mosses hugging needed for survival because they in vegetation extend also to the com- the rock faces where the winter sun dry out too rapidly. munities of mosses. In going from seldom reaches. Avoid areas where cattle are allow to high elevation, the “struc- The best place to start is to lowed to graze. Personal observa- ture” of moss populations changes locate a canyon that runs east/west, tions in the Mojave Desert in Cali- markedly: species diversity increases and has steep slopes. This canyon fornia and Nevada indicate that (more species per unit area), density should be at about 3,000 feet or where livestock grazing is presently of populations increases (mosses higher for best results. One of these occurring or has recently occurred, become easier to see because they slopes will offer a north-facing as- soil moss communities are severely are more dense), and the frequency pect, and here is where you want to damaged or absent entirely, and the of acrocarpous (having the repro- concentrate your efforts. Along a bryophytes are restricted to boul- ductive parts at the tip of a stem) north-facing slope during the win- der and rock outcrops. Recent re- populations decreases (the mosses ter and early spring months, a day search indicates that mosses take with creeping stems tend to pre- or so after a good hydrating rain- over a century to reestablish in ar- dominate).

32 FREMONTIA VOLUME 31:3, JULY 2003 RARE SPECIES FROM only from seven populations world- locality in the eastern Mojave Desert THE MOJAVE wide. Phascum hyalinotrichum is re- and another locality in southern ported from the southern Califor- Utah. This species, and at least four Perhaps oddly enough, the nia Mojave Desert, the southern others (Syntrichia bartramii, S. Mojave Desert is home to several California chaparral, the Sonoran chisosa, S. pagorum, and Didymodon globally rare or endemic species of Desert of southern Baja California, nevadensis) are of interest in that mosses and liverworts. One liver- and disjunct to the Chihuahuan males are entirely unknown. wort grows throughout the Mojave Desert of Mexico. This species ex- Desert, from Joshua Tree National hibits an unusual life history for Park to southwestern Utah, and is desert mosses: it is an annual spe- NOTABLE still undescribed to science. The cies. Because of its small size, it may CHARACTERISTICS Nevada didymodon, Didymodon be overlooked in the field. OF DESERT MOSSES nevadensis, was recently described Entosthodon planoconvexus is known from the southern Nevada Mojave, from only three populations world- The blackish, superficial, crusty and is restricted to gypsum forma- wide, one of which is in the Mojave covering often observed atop desert tions. Its distribution probably spans Desert of southern Utah, one in the soils, while inconspicuous, has great into the eastern Mojave in Califor- Sonoran Desert of Arizona (Pima ecological value to the community. nia. Despite a few reports from County), and a recent collection It consists of not only mosses, but outside deserts, Crossidium seriatum from the Isthmic Desert of north- elements from as many as four dif- remains a globally rare species that ern Egypt. ferent kingdoms. Lichens with is sporadically distributed across the Grimmia americana is known cyanobacterial symbionts fix atmo- Mojave, Sonoran, and Chihuahuan from only two populations world- spheric nitrogen. In addition to pro- deserts. It also occurs on gypsum wide, one in west Texas and the viding the primary source of nitro- and calcareous soils. other in the eastern Mojave. In the gen for the ecosystem, the desert Trichostomum sweetii appears to Mojave Desert, Pseudocrossidium crust functions in preventing soil be a southwestern endemic, known crinitum is known from only one erosion, retaining soil water, and enhancing seedling germination. Mosses of shallow soil over rocks (clockwise from top left): Syntrichia ruralis shows Desert mosses are highly vulner- hyaline awns reminiscent of those seen in Grimmia. In Syntrichia, however, the awns able to disturbance. If crushed by function as a water-repellant device: the leaves will be hydrated only in heavy rainstorms, human or animal agent (hiker, ve- and not in light sprinkles. Moss tissue will die if both wet and hot. This adaptation ensures wetting of moss leaves only if there is enough precipitation to maintain a hicle, horse), current estimates period of photosynthesis. Photograph by J. Game. • Polytrichum piliferum. Note the based on Utah communities indi- water repellant hyaline awns. Photograph by J. Shevock. • Polytrichum piliferum is cate that it will take close to two found throughout California wherever a layer of shallow soil covers rock outcrops. In centuries for them to fully reestab- desert regions, it is a plant restricted to higher elevations. The leaves of Polytrichum lish. In the Mojave Desert, where it are very thick, unlike most other mosses, which most typically have leaves one cell layer in thickness. The primary photosynthetic area is a group of photosynthetic is drier, time to full reestablishment filaments internal to the leaf. is probably longer. If you find yourself inspired by this article to learn some desert mosses, be encouraged: identification, while sometimes trying due to their small size, is easier because nearly all of the desert mosses (except those found in high mountains) fall into only two families. The dominance of the Pottiaceae and Grimmiaceae is evident to both the amateur and professional. These mosses have upright stems, grow on soil and rocks, and often have twisted leaves when dry.

Lloyd R. Stark, Department of Biological Sciences, University of Nevada, Las Ve- gas, NV 89154-4004. [email protected]

VOLUME 31:3, JULY 2003 FREMONTIA 33 THE BIOLOGY OF BRYOPHYTES, WITH SPECIAL REFERENCE TO WATER by Brent D. Mishler

he bryophytes, with more the colonization of the land by ani- Despite their diversity, phylo- Tthan 20,000 named species mals, and evolved an unparalleled genetic importance, and key roles worldwide, are the most di- diversity of size, structure, chemis- in the ecosystems of the world, study verse group of land plants (embryo- try, and function. of many aspects of the biology of phytes) except for the flowering Bryophytes contribute much to bryophytes has lagged behind that plants. The group includes three the “background” impressions of of the larger land plants, perhaps quite distinct lineages (i.e., mosses, lushness and diversity that one ex- because of their small size and the liverworts, and hornworts) of fa- periences in many habitats in Cali- few scientists specializing on the miliar species frequently encoun- fornia. But what are they, exactly, group. This is unfortunate because tered in mesic forests and along and what about them deserves to be of the intrinsic scientific value of streams, as well as a number of less brought to the “foreground” in our these plants. They have several bio- familiar species of tropical rain for- appreciation of plants? There are logical features making them par- ests, arctic tundra, and desert boul- several avenues pursued in the ar- ticularly suited to serve as study orders. The three main bryophyte lin- ticles in this issue; the direction I’ll ganisms in macroevolutionary, eages, plus a fourth, the vascular emphasize here is water. One of the population genetic, and ecological plants (tracheophytes), comprise the very most important requirements research (Mishler 1988; Shaw 1991). entirety of the monophyletic land for land plants is finding enough The plants are complicated plants (embryophytes, see figure water to get established, metabo- enough in development and ma- below), arguably one of the most lize, and reproduce. How do the ture structure to serve as model sys- important lineages to have arisen in bryophytes manage, and why? What tems for studying land plant evolu- earth’s history. They made possible general lessons can we learn from tion in general. However, they are them about how plants manage on simple enough morphologically Phylogenetic (evolutionary) tree of plants dry land? (constructed as they are from easily showing early divergence of bryophytes, First, it is necessary to go way traceable cell lineages derived from with each group (liverworts, hornworts, back in time, because the three lin- single apical cells) and genetically and mosses) as a separate lineage. This is modified from a hyperbolic tree of eages of bryophytes are modern-day (given the haploid vegetative plant green plant relationships presented on survivors of the most ancient branch- body) to be readily studied with cur- the web (http://ucjeps.berkeley.edu/ ing events in the phylogeny of the rent techniques. They are in the TreeofLife/hyperbolic.php). The tree can land plants. Fortunately, these plants great majority of cases (except for be navigated online through its thousands remain with us today, all the way some ephemeral species) observable of branches by clicking and dragging with a mouse—try it! from the spectacular radiation of the throughout the year; they are also embryophytes in the Devonian Pe- small in size, easily regenerated from riod, some 400 million years ago. As fragments, and thus easy to study in three-fourths of early land plant di- culture. versity, retaining as they do many of In a recent review (Mishler the primitive characteristics of the 2001), I asked the question: In what first land plants, bryophytes are a ways is bryophyte biology different key to understanding how the em- from that of the larger tracheo- bryophytes got started. They are also phytes? The short answer was—in the key to deciphering how plants almost every way possible! Many conquered the hostile land environ- aspects need much more study, but ment from their primitive home in what is known suggests that in gen- fresh water—habitats still occupied eral the bryophytes differ in most by relatives of the land plants, cer- ways in their biology, ecology, and tain green algae such as Chara and evolution from tracheophytes. Con- Coleochaete (Mishler and Churchill sider, for example, the following 1984, 1985; Graham 1993; Kendrick unusual features: and Crane 1997). Haploid dominance. The green,

34 FREMONTIA VOLUME 31:3, JULY 2003 vegetative part of the life-cycle in tend to have very high amounts of to follow the same historical pat- bryophytes (the “gametophyte” or morphological and physiological terns of disjunction as tracheo- gamete-bearing plant) is haploid plasticity. This may compensate for phytes, but at a lower taxonomic (i.e., it has only one set of chromo- their demonstrated low levels of ge- level. somes, like a human gamete). With- netically-based ecotypic differentia- Small stature and the occupation out the genetic benefits of domi- tion (perhaps due to haploidy). of microhabitats. Because of their nance—the masking effect of hav- Relatively slow evolutionary rates small size and lack of roots, bryo- ing two copies of each gene in a in morphology. The fossil record of phytes are in close relationship with diploid organism like our own bod- bryophytes indicates that ancient their immediate microenvironment. ies—genes acting in the gameto- forms are very similar to modern Over geological time, they may be phyte are presumably subject to ones, unlike the flowering plants less influenced by climatic change, relatively severe selection. which originated and diversified and linger in refugial habitats. Extensive phenotypic plasticity. greatly in relatively recent times. The clump as a “super-organism.” Studies have shown that bryophytes Biogeographically, bryophytes tend Many mosses and some liverworts

POIKILOHYDRY AND DESICCATION TOLERANCE: SURVIVAL STRATEGIES

he rapid equilibration of internal water content Tto the external environment is called “poikilohydry” (analogous to the term “poikilothermic” used for lizards and snakes, the so-called “cold-blooded” animals that rapidly equilibrate their internal temperature to the environment). Poikilohydry usually requires desiccation-tolerance for survival, but not necessarily. Poikilohydry is the rapid equilibration of the plant’s water content to that of the surrounding environment, while desiccation tolerance is the

ability of a plant to recover after being air-dry at the cellular level. All bryophytes have these abili- ties to some extent, unlike the larger, more complex, and endohydric tracheophytes, none of which are poikilohydric and very few of which (such as the resurrection ferns) are desiccation-tolerant.

Syntrichia ruralis (top), contrasting wet (left of twig) and dry (right of twig) plants. Photograph by B. Mishler. • Individual plants of Syntrichia ruralis (bottom), showing the widespread tendency of mosses to grow at the apex, while dying and decaying at the base. Thus most mosses live essentially forever.

VOLUME 31:3, JULY 2003 FREMONTIA 35 are essentially social organisms like a beehive. This results from the com- bination of clonal growth and external water conduction. The plants in a clump are subject to natural selection as a group. Intimate contact of each vegetative cell with the environment (due to poikilohydry—described in the sidebar on page 35), lends itself to interplant chemical communication via pheromones (Newton and Mishler 1994). In particular, let’s examine in more detail the unusual ways in which bryophytes relate to water in their environment, leftovers from the primitive past of all the early land plants. An ancestral feature of the early land plants is the con- straint imposed by the swimming sperm; there remains a need for free water for sexual reproduction to be achieved. Swimming gametes, quite feasible in the aquatic ancestral environment, are a very clumsy mechanism for land organisms (and thus were superceded in seed plants with the advent of pollen, and in land animals by internal fertilization). The fact that swimming gametes have short dispersal distances leads to frequent inbreeding in species where both sexes are borne on one plant, and leads to lack of sporophyte production in species where the sexes are borne on separate plants. Due to the difficulty of achieving fertilization, many bryophytes have evolutionarily lost func- tional sexuality. Unusually heavy re- liance on asexual reproduction has cascading effects on ecology and population genetics (Newton and Mishler 1994). Vegetative water relationships of bryophytes are even stranger by comparison to the plants you are Sphagnum sp. (top). Note the very closely spaced branches on an elongated main axis. The closeness of the branching creates capillary spaces along which water moves up used to thinking about. Unlike the the stem, an efficient alternative to the internal conducting mechanisms of the tracheophytes, they have no well- tracheophytes. • The MossCam Project (bottom). Digital video monitoring, combined developed structural mechanisms for with micro-environmental sensors, provide an automated way to continuously monitor drawing water from the soil (no the hydration status and phenology of Tortula princeps at the James Reserve. The roots), for moving water internally clump can be observed live at: www.jamesreserve.edu/mosscam/index.html along with microclimate data, a movie, and image archive. The camera has been gathering data (poorly developed conducting tis- for a year and a half, summer and winter (when the mosses are most active). Photograph sue), or for holding water inside by S. Lubin. (leaves are only one to two cells thick

36 FREMONTIA VOLUME 31:3, JULY 2003 Examples of some desiccation-tolerant mosses (clockwise from top left): Syntrichia ruralis often goes through several cycles of drying and wetting, even in a single day, without apparent damage to the plant. • Grimmia torquata. In a genus especially adapted to growth on dry rocks, G. torquata is an exception. Its dense cushions are found on rock overhangs where protected from direct rainfall. While it has hyaline awns, like most of the rest of the genus, those awns are very reduced in size. It is also a unique member of the genus in having leaves that dry in a disorderly manner (crispate). • Racomitrium lanuginosum is another member of the Grimmiaceae, but members of this genus grow primarily on very sunny rocks in humid coastal areas. In Alaska and high arctic, R. lanuginosum often grows in such large colonies as to be identified from as far as one mile away. • Racomitrium heterostichum is our most common member of the genus Racomitrium, and is often found covering cliffs and roadcuts, especially in northwestern California. In its typical habitat it may be dry and dormant for ninety percent of the year. • Orthotrichum consimile belongs to one of the largest genera of mosses in California. Members of this genus are found growing on both rocks and trees, and obtain essentially all of their water from rapid absorption of droplets on the above-ground plant. Like all mosses, species of Orthotrichum have no root system penetrating the substratum.

VOLUME 31:3, JULY 2003 FREMONTIA 37 with thin cuticles). Instead, bryo- genus Sphagnum, which acidifies its about these remarkable little plants. phytes have about the same water environment by trading hydrogen We need your help to understand relationships as a piece of paper sit- ions for other cations. Mosses (along them better. ting on a rock—if it rains it gets wet with lichens which are also fast, if it stops raining it gets dry fast. poikilohydric) are quite sensitive to In a recent study, Oliver, Tuba, water and air quality and thus are REFERENCES and Mishler (2000) showed that veg- important bioindicators. They can etative desiccation tolerance was also be quite choosy about their Graham, L.E. 1993. The origin of land primitively present in the land plants habitats, each species growing on plants. Wiley. New York, NY. (as seen today in all bryophytes), particular bark, rock, or soil types. Kenrick, P., and P.R. Crane. 1997. The but was then lost in the evolution of We know little about the actual origin and early diversification of land tracheophytes. The initial evolution water budget of these poikilohydric plants: A cladistic study. Smithsonian Institution Press. Washington, DC. of vegetative desiccation-tolerance plants, i.e., how often they are wet, Mishler, B.D. 1988. Reproductive was a crucial step required for the for how long, and at what fre- ecology of bryophytes. In Plant recolonization of the land, but that quency. To know this would reproductive ecology, ed. Lovett Doust, tolerance came at a cost, since meta- quire constant monitoring of popu- J. and L. Lovett Doust, 285-306. bolic rates are lower in tolerant lations over years, which is impos- Oxford University Press. London. plants as compared to plants that sible for someone to do first-hand. Mishler, B.D. 2001. Review of The bi- don’t maintain costly mechanisms To begin to approach this ques- ology of bryophytes edited by Shaw and for tolerance. Thus, the loss of tol- tion, Mike Hamilton and Sheri Goffinet, American Journal of Botany erance might have been favored Lubin (of the James Reserve in the 88:2129-2131. along with the internalization of San Jacinto Mountains in South- Mishler, B.D., and S.P. Churchill. water relationships that happened ern California) and I initiated a re- 1984. A cladistic approach to the phylogeny of the “bryophytes’. as the vascular plants became more mote-sensing project using video Brittonia 36:406–424. complex and able to control their cameras hard-wired to the Internet Mishler, B.D. and S.P. Churchill. internal water balance. (dubbed the “MossCam” project; 1985. Transition to a land flora: However, at least two indepen- see photograph on page 36 and Phylogenetic relationships of the dent evolutions (or re-evolutions) www.jamesreserve.edu/mosscam/). green algae and bryophytes. Cladis- of desiccation-tolerance occurred in We are monitoring clumps of the tics 1:305–328. Selaginella and in some ferns. diverse moss genus Syntrichia, look- Mishler, B.D., L.A. Lewis, M.A. Within the flowering plants, at least ing at periodicity of hydration, and Buchheim, K.S. Renzaglia, D.J. eight independent cases of evolu- seasonality of reproduction. In ad- Garbary, C.F. Delwiche, F.W. tion (or re-evolution) of vegetative dition to its serious scientific side, Zechman, T.S. Kantz, and R.L. Chapman. 1994. Phylogenetic rela- desiccation-tolerance occurred. The this project also provides an excit- tionships of the “green algae” and time scale and mechanisms of des- ing website where one can watch “bryophytes.” Annals of the Missouri iccation and rehydration are differ- moss grow in real time! Botanical Garden 81:451–483. ent each time the general pheno- Many other aspects of the biol- Newton, A.E., and B.D. Mishler. 1994. type was re-evolved. Deciphering ogy of bryophytes are in need of The evolutionary significance of the physiological mechanisms and study and could yield important re- asexual reproduction in mosses. genes behind these complex phe- sults. Many of the observations Journal of the Hattori Botanical Labo- notypes is an exciting area of cur- needed are very low-tech, and could ratory 76:127–145. rent research with both intellectual be carried out by any general bota- Oliver, M.J., Z. Tuba, and B.D. and economic applications (for nist. We need to know much more Mishler. 2000. The evolution of vegetative desiccation tolerance in more information see the “Plants about simple demography (i.e., how land plants. Plant Ecology 151:85– Without Water” website at: http:// new plants are established and how 100. ucjeps.herb.berkeley.edu/bryolab/pww). they die), phenology (the annual Shaw, A.J. 1991. Ecological genetics, Poikilohydry defines many as- timing of production of gametan- evolutionary constraints, and the pects of bryophyte ecology. Since gia and sporophytes, the diploid systematics of bryophytes. Advances water is moving back and forth di- phase), distribution patterns (both in Bryology 4:29–74. rectly across the membrane of the geographic and ecological), use by photosynthetic cell, these mem- animals, and many other topics that Brent D. Mishler, University Herbarium, branes are in unusually intimate are accessible and would lend them- 1001 Valley Life Sciences Building, No. contact with their immediate mi- selves to simple field research. Like 2465, University of California, Berkeley, croenvironment. They are famous the other authors in this special is- CA 94720-2465. bmishler@socrates. for ion exchange, most notably the sue, I urge you to find out more berkeley.edu

38 FREMONTIA VOLUME 31:3, JULY 2003 FROM THE ARCHIVES: BRYOPHYTES & LICHENS

his is not the first Fremontia fungus (not a plant; provides the Tissue with articles on non- scientific name for the lichen) asso- vascular plants. As men- ciated with an alga (photosynthe- tioned on page 12, Jim Shevock sizes within the structure formed wrote an article about bryophytes by the fungus and provides energy that appeared in the April 1998 to the organism). See the July 1984 issue of Fremontia (Volume 26(2): issue of Fremontia (Volume 12(2): 3–8). You may wish to read this 21–22) for an article by Janet H. article as an introduction to the cur- Wood (then studying lichens for rent issue, as Jim describes the basic her Master’s thesis) on how lace characteristics of byrophytes in gen- lichen (Ramalina menziesii) is used eral, followed by a more in-depth to indicate air pollution levels. Lace discussion of mosses in particular, lichen is often found draping from including distributions and notes on oak trees. collecting them. The section on The April 1993 issue (Volume field guides in this article is some- 22(2):3–12) has an excellent article what dated; see this issue’s Notes on lichens written by Wayne P. and Comments (page 40) for Jim’s Armstrong (then biology professor current comments under “Building at Palomar College in San Marcos) a Bryophyte Library.” and Jamie L. Platt (then student at The California Native Plant California State University, San Society is also interested in lichens, Marcos), where the authors fully which are not technically plants, but describe the relationship between rather are organisms consisting of a the fungus and alga that together Cover of the April 1998 issue of Fre- form the lichen. The authors also montia, showing the trunk of a large interior live oak (Quercus wislizenii) Cover of the July 1999 issue of Fremontia, discuss the structure of a lichen and covered with the mosses Dendroalsia with pixie-cup lichen (Cladonia asahinae), the three growth forms (with lovely abietina and Homalothecium sp. (the shown growing on the base of a tree in photographs), along with the role original photograph by M. Bourell). Carmel (the original photograph by S. Sharnoff). of lichens in succession, uses of lichens by natives and explorers, and late wife Sylvia D. Sharnoff’s spec- the diminished range of lichens due tacular lichen photography, and was to air pollution. a prelude to their book (with I.M. Stephen Sharnoff’s “Lichens: A Brodo, lichenologist), Lichens of Different Window on California’s North America. A book review of Diversity” appears in the July 1999 this volume appears on page 41 issue (Volume 27(3):10–13). This along with two examples of Sharnoff article is graced with Steve’s and his lichen photography.

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ack issues of Fremontia are available for sale from the CNPS BOffice, 2707 K Street, Suite 1, Sacramento, CA 95816; phone (916) 447-2677. Issues for Volume 28 and later (2000-present): $5 each or $10 for three. Issues before Volume 28: $2.50 each or $6 for three. Double issues priced as two single issues; shipping costs determined upon order placement.

VOLUME 31:3, JULY 2003 FREMONTIA 39 NOTES AND COMMENTS

BUILDING A BASIC bia. Royal British Columbia Press. fore, it can be viewed as a reference BRYOPHYTE LIBRARY Victoria, BC. for someone that has already had a A nice paperback guide with black general bryophyte course or desires and white illustrations. Useful for more information after reading either The bryological literature is wide- moss identifications along the coast Crum (2001) or Schofield (1985). spread, residing among numerous redwood belt of northwest California. journals and other books. Some bryo- Schofield, W.B. 2002. Field Guide floras and identification manuals are to Liverwort Genera of Pacific North BRYOLOGICAL currently out of print. However, there America. Global Forest Society, San INTERNET are several key works that are avail- Francisco and University of Washing- able and quite useful to begin a study ton Press. Seattle, WA. RESOURCES of California bryophytes. These pub- A really nice paperback with full- lications are organized under the titles page illustrations of each genus. Avail- The following is a brief list of use- of bryofloras, field guides, and text- able through Washington University ful Internet sites for both the begin- books. Press [www.washington.edu/press]. ning and professional bryologist. American Bryological and Bryofloras Textbooks Lichenological Society (ABLS), Flowers, Seville 1973. Mosses: Utah Crum, Howard H. 2001. Structural www.unomaha.edu/~abls. and the West. Brigham Young Univer- Diversity of Bryophytes. University of The ABLS publishes two quarterly sity. Provo, UT. Michigan Herbarium [phone order bryological journals. The more tech- This book contains excellent illus- 734-764-2407]. nical of the two is The Bryologist and trations and has recently been re- This is the last work of the late the other is Evansia. The ABLS home printed by Blackburn Press [www. Howard Crum and it is a gem. It is page provides numerous links to other blackburnpress.com]. It is very useful for well written and illustrated. bryological websites around the world. the identification of mosses from the Schofield, W.B. 1985 (reprinted Other bryological societies and jour- more arid regions of California, espe- 2001). Introduction to Bryology. Black- nals can be accessed through this site. cially the deserts and southern Cali- burn Press. Caldwell, NJ. Missouri Botanical Garden—Bryol- fornia. This is an excellent textbook (suit- ogy Lab (MO), www.mobot.org/ Lawton, Elva 1971. Mosses of the able too for college courses) that is also MOBOT/.research/links5.shtml. Pacific Northwest. Hattori Botanical well illustrated. This page provides links to a wide Laboratory. Miyazaki-ken, Japan. Shaw, J. and B. Goffinet 2000. Bryo- variety of bryological institutions This book is available in a paper- phyte biology. Cambridge University throughout the world. Bryology at back edition from the Hattori Botani- Press. Oxford, UK. MO has several floristic projects un- cal Laboratory [[email protected]. This paperback edited by Shaw and derway and the herbarium has over ne.jp]. It is very useful for the coastal Goffinet contains 13 chapters written 300,000 bryophyte collections. In ad- counties of northern California and by leaders in their bryological field of dition, a complete list of bryophyte mountainous portions of the state. Il- expertise. It is a bit more technical names can be accessed electronically lustrations are provided but not as than either Crum or Schofield. There- at this site. detailed nor of the same quality as New York Botanical Garden (NY), those from Mosses: Utah and the West. www.nybg.org/bsci/hcol/bryo/bryology_ center.html. BRYOPHYTE WORKSHOPS The NY site provides access to Field guides many bryological sites and useful in- February 21-22, 2004 Malcolm, B. and N. Malcolm, 2000. formation. The North American Mosses and Other Bryophytes: An Illus- Jepson Bryophyte Workshop bryophyte specimens at NY (the trated Glossary. Micro-optics Press. For more information, Nation’s largest herbarium) can be New Zealand. call (510) 643-7008, or see searched on line. While not a field guide, this illus- http://ucjeps.berkeley.edu/jepwkshp.html IUCN World Red List of Bryo- trated glossary contains 970 color March 20-23, 2004 phytes, www.artdata.slu.se/guest/ images to provide a visual aid to defi- SSBryo/Bryolist.htm. nition of bryophyte features and ter- SO BE FREE, West Coast Foray This section of the IUCN is work- minology. This book is essential when Sponsored by Brent Mishler, ing to save the worlds most endan- University Herbarium, UC Berkeley using bryophyte keys for identification gered mosses, liverworts, and horn- For more information, see purposes and is highly recommended. http://usceps.herb.berkeley.edu/bryolab/ worts. Two California liverworts Schofield, W.B. 1969 (reprinted). trips/sobefreel.html (Geothallus tuberosus and Sphaerocarpus Some Common Mosses of British Colum- drewei) are on the World Red List.

40 FREMONTIA VOLUME 31:3, JULY 2003 BOOK REVIEWS

Moss Gardening: Including Li- me how to get started in learning and of their importance. She clearly chens, Liverworts, and Other Min- about bryophytes and what books to has a cultural kinship to the mosses iatures, by George Schenk. 1997. 262 buy. A popular, non-technical, easy- which, in many ways, makes this book pages, with 97 color photos. Timber to-read yet scientifically accurate book unique. It reads like a natural history, Press (www.timberpress.com). Portland, has finally crossed my desk. but one with a considerably more per- OR. Price $34.95, hardcover. Robin Kimmerer’s book is a great sonal tone to it. Despite its non-tech- Appearing in fossils 400 million place to start for those unfamiliar with nical approach, however, the book years old and totaling 15,000 species mosses. Kimmerer is a biologist (as- provides information about general from the Arctic to the Antarctic, sociate professor) and a Native Ameri- ecology as well as examples of scien- mosses tell an ancient success story of can woman who shares her experi- tific principles. longevity and dispersion. North ences of how the mosses speak to us Each of the book’s short essays pre- America alone supports 1,200 of them. George Schenk’s beautiful book is per- The lichen Cladonia carneola on a rotting log. Photograph by S.D. Sharnoff. haps the first to cover the whole gamut of moss gardening. Moss Gardening is sure to become a classic, not only because it covers “almost as much about the mechanics of moss gardening as the Kama Sutra does about dancing,” but also for the author’s entertaining writing style. The book covers primitive minia- tures (or cryptogams) which reproduce by spores: mosses, lichens, liverworts, lycopodiums, and selaginellas. The information on propagating, cultivating, and transplanting is ap- plicable to gardens of practically any climate. Landscaping recommenda- tions are offered for use in alpine and rock gardens, with flowering plants in borders, as a lawn substitute, and for use in miniature gardens. George Schenk retired from his landscape and nursery business and now designs and maintains gardens for friends in Seattle, North Van- couver, Auckland, and Manila. He authored of three other horticultural books. Anonymous Timber Press

Gathering Moss, A Natural and Cultural History of the Mosses, by Robin Wall Kimmerer. 2003. Oregon State University Press. Corvallis, OR. Orders processed through collabora- tion with University of Arizona Press. Tucson, AZ. 168 pages. Price $17.95, hardcover. The mosses (and other bryophytes) are a remarkable group of plants, and many botanists and plant enthusiasts have yet to be exposed to their diminu- tive beauty. Many people have asked

VOLUME 31:3, JULY 2003 FREMONTIA 41 sents a particular story and lesson about the role mosses play in our world. Here are a few passages to give you the flavor of Kimmerer’s prose and how she brings us into the world of mosses. “. . . Learning to see mosses is more like listening than looking. A cursory glance will not do it. Straining to hear a far- away voice or catch a nuance in the quiet subtext of a conversation requires atten- tiveness, a filtering of all the noise, to catch the music. Mosses are not elevator music; they are the intertwined threads of a Beethoven quartet. You can look at the mosses the way you listen deeply to running water over rocks. The soothing sound of a stream has many voices, the soothing green of mosses likewise.” “. . . At the scale of a moss, walking through the woods as a six-foot human is a lot like flying over the continent at 32,000 feet. So far above the ground, and on our way to somewhere else, we run the risk of missing an entire realm which lies at our feet. Every day we pass over them without seeing. Mosses and other small beings issue an invitation to dwell for a time right at the limits of ordinary per- ception. All it requires of us is attentive- ness. Look in a certain way and a whole new world can be revealed.” The closing passage of the Preface sets the stage for all of the essays that follow. “. . . In indigenous ways of knowing, we say that a thing can not be understood until it is known by all four aspects of our being; mind, body, emotion, and spirit. The scientific way of knowing re- lies only on the empirical information from the world, gathered by the body and in- terpreted by the mind. In order to tell the The lichen Lecanora caesiorubella on oak bark. Photograph by S.D. Sharnoff. mosses’ story I need both approaches, ob- jective and subjective. These essays inten- pleted it, your next walk in a forest may tographs of several hundreds of lichen tionally give voice to both ways of know- very well seem like your first! species, useful to the specialist, the ing, letting matter and spirit walk com- James R. Shevock amateur, and the natural historian panionably side by side. And sometimes Department of Botany with general interests. The second even dance.” California Academy of Sciences edition of this opus, published as two As an ecologist, Kimmerer has stud- volumes in 1995, also included the ied mosses in the field for countless Lichens of North America, by well-elaborated keys known from hours and through this association I.M. Brodo, S.D. Sharnoff, and S. Wirth’s Flechtenflora, and additional their secrets have been revealed to her. Sharnoff. 2001. Yale University Press, photographs of more species. She now shares them freely with the New Haven and London. 795 pp. None, including myself, would have reader. It is a rare event for me to read Price $ 69.95, hardcover. believed it possible to match or sur- a botany book from cover to cover With Die Flechten Baden-Württem- pass the superb quality of Volkmar without jumping between chapters. bergs, fifteen years ago Volkmar Wirth Wirth’s books. Yet, here we are with This is just such a book—unlike any- set the standard for what would ever another book for which it is difficult, thing I have read in a long time. I since have been the dream for a lichen if not impossible, to find superlatives, found it to be a delight and I suspect monographer: a book in high print and which not only matches, but sur- you will too. And once you’ve com- quality, filled with first class color pho- passes any other similar book pub-

42 FREMONTIA VOLUME 31:3, JULY 2003 lished in this area: Brodo et al’s Lichens dict that this book will have a signifi- is the late Sylvia Sharnoff, S. Sharnoff of North America. When I first saw this cant impact on, and raise the numbers is Stephen Sharnoff] range among the book, I simply couldn’t believe that of, students of lichenology. And if best nature and plant photographers something as wonderful as this really there is one most important achieve- in the world. Lichens of North America existed. But it does, and it is not only ment of any scientific book, it is the is definitely their masterpiece. affordable, in fact, the price of this attraction of students to the subject. Robert Lücking book is quiet incredible [but note that Irwin Brodo has marked lichenology Lichen Collection Manager at time of publication of this Fremontia in North America and beyond for de- Field Museum of Natural History issue, less than 300 copies remained], cades. The Sharnoffs [S.D. Sharnoff Chicago considering that it features nearly 1,000 color photographs of absolutely superb quality. CONTRIBUTORS (Cont’d from back cover) One could certainly criticize a few minor weak points of this opus. For example, many line drawings are not both in western North America and plants in the Sierra Nevada. He is re- originals but reproduced from other in neo- and paleotropical regions. His search coordinator for the National books. Or, compared to Wirth’s bryophyte collection, at over 106,000 Park Service at the Californian Coop- books, comparatively few crustose specimens, is one of the largest in the erative Ecosystem Studies Unit. lichen species are represented. Also, world, and forms the nucleus of the Lloyd Stark, PhD, is an assistant pro- in most genera, the keys to species are University Herbarium bryophyte col- fessor at the University of Nevada, Las not complete but rather represent a lection. selection of common North Ameri- Vegas, Department of Biological Sci- can taxa. James R. Shevock, research associ- ences. His research explores how male However, this is not the moment to ate with the Department of Botany, and female desert mosses respond to criticize. In fact, this book is beyond California Academy of Sciences, and stresses of extreme temperatures and criticism, because its primary goal is the University Herbarium, UC Ber- desiccation, and also tracks the distri- not to be a complete lichen flora of keley, is one of the foremost authori- butions of rare aridland species of North America. This book is differ- ties on both bryophytes and vascular mosses. ent. This book is probably the most important work ever published on lichens, because it addresses the widest audience possible and will, without CLASSIFIED ADS doubt, attract hundreds, if not thousands of students and nature lovers to this fascinating and important group Classified ad rate: $1.00 per word, Botanical prints, note cards, postcards, minimum $15; payment in advance. Plants of the Lewis and Clark Expedition of organisms. And, there are two sig- Address advertising inquiries and copy to: notecards, books, fruit crate labels, hand nificant advantages over Wirth’s CNPS, 2707 K Street, Suite 1, Sacra- towels, and T-shirts focusing on books. First, the introductory part of mento, CA 95816-5113. (916) 447-2677 California native plants from Sierra Lichens of North America is much more or fax (916) 447-2727. Nature Prints. Animal puppets, too. Visit www.sierranatureprints.com extensive, covering all aspects of lichen PUBLICATIONS biology and their potential uses, and NURSERIES AND SEEDS Flora & Fauna Books, 121 First Avenue second (trivial but not less important!), Telos Rare Bulbs. Bulbs for your gar- this book is in English, and hence it is South, Seattle WA 98104, Tel. (206) 623-4727, Fax (206) 623-2001, ffbooks den, restoration projects, landscaping. accessible to anybody with a basic @blarg.net, Specializing in Botany, Many Calif. native species, including Calochortus, Fritillaria, Brodiaea rela- knowledge of that language. Gardening, Birding, and Ecology, both tives, Erythronium. Catalog $3.00. Free I have already experienced the im- new and out-of-print. We carry a large shipping in USA. P.O. Box 4978, Arcata, pact of Lichens of North America on sev- inventory of floras, keys, and field guides CA 95518. www.telosrarebulbs.com. eral occasions, most recently during a for the west coast and worldwide. A large selection of our inventory is now ECOLOGICAL RESTORATION visit to an herbarium (UNAM) in available on the web: www.abebooks.com/ Mexico, where Marusa Herrera Cam- home/FFBOOK/. Native California grassland and desert pos has successfully established a ecological restoration standards and lichenological working group, and all ART costs, plus pictures showing results. www.ecoseeds. com/standards.html. the students, regardless of whether California wildflower prints & free they could possibly afford it or not, wildflower screensavers. Visit www. SERVICES unanimously stated: “I want that wildflowergreetings.com or call Alice (877) 432-2999 toll-free. Nature landscape design. Landscape book!” It is certainly no overstatement Design that celebrates the rich heritage to say that the Lichens of North America, Notecards, Prints, and Originals. Visit of California’s native flora. Duber Land- from now on, will be the bible for www.VorobikBotanicArt.com. 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VOLUME 31:3, JULY 2003 FREMONTIA 43 FROM THE EDITOR

re you ready to appreciate the Next, Jim Shevock provides an over- into collecting and cataloguing those A small and wonderful? Pre- view of the mosses that occur in Cali- from the environs of his Santa Cruz pare yourself to spend some fornia, where they might be found, home. field time on your knees, or pressed and some of their characteristics. He Think mosses are only from wet ar- against tree trunk or rock, after read- then teases us with names of several eas? Wonder how mosses survive con- ing this issue and becoming enthused mosses that should be in California, tinual wetting and drying, without real to learn more about mosses, liver- but that have yet to be discovered, roots? Consider the paper by Lloyd worts, and hornworts. combined with the information that Stark, who studies mosses of desert The issue begins conversationally: new species of mosses are found in our regions, or that of Brent Mishler, who Dan Norris provides general informa- state every year! is very interested in how mosses uti- tion about these small green plants in If you quake with the trepidation of lize the water in their environment. a series of questions that he has heard a novice, read Ken Kellman’s article. Read, enjoy, research more, but do get throughout his career as bryology pro- He, like most bryologists, started with down on your knees to glory in the fessor. How better to learn about a the vascular plants (wildflowers, in greatness of these tiny green plants. group than to read the answers to the fact), but once he learned a bit about Linda Ann Vorobik most frequently asked questions? mosses he was irretrievably hooked Editor CONTRIBUTORS

John Game works in molecular genetics for the Lawrence Berkeley National Laboratory, with degrees in botany from Oxford, England. He is a research associate at the UC/

Jepson Herbaria, an active California Native Plant Society

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2707 K Street, Suite 1 Suite Street, K 2707 Sacramento, CA 95816-5113 CA Sacramento, member, as well as an enthusiastic plant photographer with Society Plant Native California many images published in books, magazines, and on the internet. Martin Hutten received a graduate degree in forest science at Oregon State University studying relationships between mycorrhizae and trees, and has spent several years conducting an inventory of lichens, mosses, and liverworts for Olympic National Park. His excellent photographs populate this entire issue of Fremontia. Kenneth Kellman is an amateur botanist who has been studying bryophytes since 1995. He is currently working on “A Catalog of the Liverworts and Hornworts of Santa Cruz County”, and is starting collections for a bryophyte flora of Monterey County. Brent Mishler, PhD, is director of the Jepson and Uni- versity Herbaria at UC Berkeley, as well as a professor in the Department of Integrative Biology, where he teaches systematics and plant diversity. His research interests are the systematics, evolution, and ecology of bryophytes, as well as the phylogeny of green plants and the theory of systematics. Daniel Norris, PhD, is research botanist at the University

California, Berkeley, with extensive bryological experience

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FREMONTIA EDITORIAL ADVISORY BOARD Ann Bradley, Travis Columbus, Susan D’Alcamo-Potter, Ellen Dean, Kathleen Dickey, Phyllis M. Faber, Bart O’Brien, John Sawyer, Jim Shevock, Teresa Sholars, Nevin Smith, Dieter Wilken, John Willoughby, Darrell Wright 44 FREMONTIA VOLUME 31:3, JULY 2003