Fremontia Vol. 45, NOS 1 and 2, Page 30

Home , Calicium, Comanche Point, Eriophyllum lanatum, Pseudotsuga macrocarpa, Ramalina menziesii, Santa Ana Mountains

$10.00 (Free to Members)

VOL. 45, NOS.1&2 • JUNE 2017 FREMONTIA JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

LICHENSLICHENS ANDAND BRYOPHYTESBRYOPHYTES TEJONTEJON RANCHRANCH REVISITEDREVISITED BIGCONEBIGCONE DOUGLAS-FIRDOUGLAS-FIR VOL. 44, NO. 3, DECEMBER 2016 FREMONTIA VEGETVEGETATIONATION MAPPINGMAPPING ANDAND RE-MAPPINGRE-MAPPING

V45_2_cover.pmd 1 6/9/17, 6:51 AM CALIFORNIA NATIVE PLANT SOCIETY CNPS, 2707 K Street, Suite 1; Sacramento, CA 95816-5130 FREMONTIA Phone: (916) 447-2677 Fax: (916) 447-2727 Web site: www.cnps.org Email: [email protected] VOL. 45, NOS. 1&2, JUNE 2017 MEMBERSHIP Copyright © 2017 Members receive many benefits, including subscriptions to Fremontia and California Native Plant Society the CNPS Bulletin. Membership form is on inside back cover. Mariposa Lily ...... $1,500 Family or Group ...... $75 Michael Kauffmann, Editor Benefactor ...... $600 International or Library ...... $75 Patron ...... $300 Individual ...... $45 Beth Hansen-Winter, Designer Plant Lover ...... $100 Student/Retired/Limited Income . $25 CORPORATE/ORGANIZATIONAL ISSN 0092-1793 10+ Employees ...... $2,500 4-6 Employees ...... $500 7-10 Employees ...... $1,000 1-3 Employees ...... $150

california Native STAFF & CONTRACTORS Plant Society Dan Gluesenkamp: Executive Director Milo Baker: Leia Giambastiani, Sarah Chris Brown: Admin Assistant Gordon Protecting California’s Native Flora Jennifer Buck-Diaz: Vegetation Ecologist Mojave Desert: Timothy Thomas Since 1965 Catherine Curley: Assistant Botanist Monterey Bay: Christopher Hauser Julie Evens: Vegetation Program Dir. Mount Lassen: Woody Elliot The views expressed by authors do not Stacey Flowerdew: Membership & Napa Valley: Henni Cohen necessarily reflect established policy or Development North Coast: Carol Ralph procedure of CNPS. Vern Goehring: Legislative Analyst North San Joaquin: Jim Brugger Kaitlyn Green: Assistant Rare Plant Orange County: Dan Songster Botanist Redbud: Denise Della Santina Laureen Jenson: Accounting & HR Riverside/San Bernardino: Katie Barrows Michael Kauffmann: Fremontia Editor Sacramento Valley: John Hunter David Magney: Rare Plant Program Mgr. San Diego: Bobbie Stephenson North Coast Shasta Statewide Chapters Mark Naftzger: Webmaster San Gabriel Mtns.: Orchid Black Liv O’Keeffe: Marketing and Comms. Dir. San Luis Obispo: Bill Waycott

Bryophyte Chapter Jaime Ratchford: Assoc. Veg. Ecologist Sanhedrin: Jennifer Riddell

Mt. Lassen Becky Reilly: Events Coordinator Santa Clara Valley: Stephen Rosenthall Kendra Sikes: Vegetation Ecologist Santa Cruz County: Deanna Giuliano Tahoe Aaron Sims: Rare Plant Botanist Sequoia: Vacant Sanhedrin Redbud Sacramento Dorothy Greg Suba: Conservation Program Dir. Shasta: Ken Kilborn Milo El Dorado King Young Napa Baker Valley Karen Whitestone: Conservation Analyst Sierra Foothills: Bob Dean Willis Linn Jepson Brock Wimberley: Ops & Finance Dir. South Coast: David Berman Sierra Foothills Marin East Tahoe: Brett Hall Bay North TheSan Joaquin bright orange splash of lichen at the peak of this outcrop tells us something—there’s nitrogenWillis input L. Jepson: at that Stephen spot. GoetzIn this case, the Yerba Buena Santa CHAPTER COUNCIL—CHAPTERS Clarasource is birds perching-and pooping-on the rock’s summit. Photographs by Shelly Benson unless otherwise noted. Valley & DELEGATES Yerba Buena: Ellen Edelson Santa Cruz County Sequoia Bristlecone Marty Foltyn: Chair Monterey Larry Levine: Vice Chair 2017 BOARD OF DIRECTORS Bay SEEINGAlta Peak LICHENS. . . AND WHAT THEY TELL US Judy Fenerty: Secretary Steve Hartman: President ABOUT OURAlta Peak: ENVIRONMENT Melanie Keeley Bill Waycott: Vice President San Luis Obispo Kern Baja: César GarcÌa Valderrama Gabi McLean: Treasurer Mojave Bristlecone:by Shelly Katie Benson Quinlan Carolyn Longstreth: Secretary Bryophyte: Paul Wilson Julie Clark DeBlasio: Director Channel Islands San Gabriel Mtns. Channel Islands: Andrea Adams-Morden Brett Hall: Director Riverside – San Bernardino Dorothy King Young: Nancy Morin David Pryor: Chapter Council Rep. Los Angeles – Orange ichensSanta are Monica found Mtns. County all around us, the object,East a Bay: recognition Barbara Leitner link is made can’tGordon believe Leppig: that Director they had been walk- El Dorado: Alice Cantelow Cari Porter: Director growing in nearly every Sanhabitat Diego and one cannot help but see it in ing blindly past this diverse group and onSouth any Coast stable substrate. Yet, one’s surroundings.Kern County: Rich As Spjut a lichenolo- ofJean organisms Robertson: for Director so long. Seeing the they are often overlooked be- gist, myLA/Santa mission Monica is to Mtns.: help Snowdy people Dodson amazingCris Sarabia: colors, Director shapes, and textures Marin: Charlotte Torgovitsky Vince Scheidt: Director Lcause many people are unsureBaja of “see” lichens. When I lead nature of lichen can give great enjoyment. California walks, I explain the basics about what lichens are. I once heard some- MATERIALS FORBut PUBLICATION that’s not all—lichens are widely one speak on the topic of how words lichens CNPSand pointmembers out and examples others are welcome of toused contribute as indicators materials forof airpublication quality, in cli- affect our ability to see. The speaker’s the differentFremontia lichen. See the growth inside backforms: cover for mate,submission and instructions. old growth forests; once point was that an object right under foliose, fruticose, and crustose. Al- you learn to see lichens you can Staff and board listings are as of June 2017. your nose could go unnoticed until most immediately, peoplePrinted bybegin Modern to Litho: discoverwww.modernlitho.com the stories that they tell you have a word for it. By naming observe lichens everywhere. They about our environment.

FREMONTIA2 FREMONTIA VOL.VOL. 45,44, NOS. NO. 3, 1 DECEMBER& 2, JUNE 2017 2016

V45_2_cover.pmdV45_2_book.pmd 2 6/9/17, 6:516:52 AM CONTENTS

SEEING LICHENS…AND WHAT THEY TELL US ABOUT OUR ENVIRONMENT by Shelly Benson ...... 2 Lichens display an amazing variety of colors, shapes, and textures. But that’s not all— lichens are widely used as indicators of air quality, climate, and old-growth forest health.

LICHEN CONSERVATION IS COMPLICATED by Rikke Reese Næsborg ...... 8 Like other rare, endangered, or threatened organisms lichens need protection, but management decisions are challenged by the lack of knowledge about lichen distribution and ecology.

A BIT ABOUT BRYOPHYTES by Marie Antoine ...... 12 Lichens and bryophytes co-occur in various ecological niches. “Bryophyte” is a general term referring to mosses, liverworts, and hornworts. These tiny plants are often overlooked, but close observation reveals a fascinating miniature world with an intricacy of form and diversity of function.

TIMES THEY ARE A CHANGING: VEGETATION MAPPING AND RE- MAPPING EFFORTS ACROSS CALIFORNIA by Todd Keeler-Wolf, Rachelle Boul, Jennifer Buck-Diaz, and John Menke ...... 18 This article explores the value of repeat vegetation mapping while explaining the careful steps that must be taken to guarantee highest value for change detection.

TEJON RANCH—LAND OF CONTRAST, BOTANICAL RICHNESS, AND ONGOING DISCOVERY by C. Ellery Mayence, Nick Jensen, Neal Kramer, Laura Pavliscak, and Michael D. White ...... 25 Tejon Ranch, an amalgamation of numerous iconic California landscapes, is a hub of biodiversity conservation, environmental education, and scientific research in the Tehachapi Region.

BIGCONE DOUGLAS-FIR MAPPING AND INVENTORYING IN THE ANGELES NATIONAL FOREST by Michael Kauffmann, Julie Evens, and Jaime Ratchford ...... 30 The California Native Plant Society, in collaboration with the US Forest Service, assessed the extent and status of the California endemic bigcone Douglas-fir (Pseudotsuga macrocarpa) in the Angeles National Forest.

BOOK REVIEWS ...... 42

On the cover: Lichens glow at sunrise, decorating the upper redwood (Sequoia sempervirens) canopy in Redwood National and State Parks. Photo by Marie Antoine.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 1

V45_2_book.pmd 1 6/9/17, 6:52 AM The bright orange splash of lichen at the peak of this outcrop tells us something—there’s nitrogen input at that spot. In this case, the source is birds perching-and pooping-on the rock’s summit. Photographs by Shelly Benson unless otherwise noted. SEEING LICHENS. . . AND WHAT THEY TELL US ABOUT OUR ENVIRONMENT by Shelly Benson

ichens are found all around us, the object, a recognition link is made can’t believe that they had been walk- growing in nearly every habitat and one cannot help but see it in ing blindly past this diverse group and on any stable substrate. Yet, one’s surroundings. As a lichenolo- of organisms for so long. Seeing the they are often overlooked be- gist, my mission is to help people amazing colors, shapes, and textures Lcause many people are unsure of “see” lichens. When I lead nature of lichen can give great enjoyment. what lichens are. I once heard some- walks, I explain the basics about But that’s not all—lichens are widely one speak on the topic of how words lichens and point out examples of used as indicators of air quality, cli- affect our ability to see. The speaker’s the different lichen growth forms: mate, and old growth forests; once point was that an object right under foliose, fruticose, and crustose. Al- you learn to see lichens you can your nose could go unnoticed until most immediately, people begin to discover the stories that they tell you have a word for it. By naming observe lichens everywhere. They about our environment.

2 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 2 6/9/17, 6:52 AM WHAT IS A LICHEN? NEWS FLASH: Two’s Company, Three’s a Lichen? It is a composite organism made up of fungus and a photosynthesiz- ecent scientific findings from Spribille et al. (2016) change our ing partner (photo-partner). Last Runderstanding of the lichen relationship. For over 140 years we summer a team of lichenologists an- thought that lichens were made of a fungus and an alga or nounced their discovery that many cyanobacteria. Spribille’s team discovered that many lichen species lichens contain two types of fungi contain two types of fungi. The first type (the one we already knew (see NEWS FLASH). The photo- about) is typically in the group ascomycetes—the sac fungi. The partner can be either a green alga or previously unseen fungal partner is a newly discovered lineage of a cyanobacteria. In a few lichens, yeast in the group basidiomycetes—the club fungi, which include both types of photo-partners are gilled mushrooms. These yeasts appear to be responsible for produc- present. This oddity is interesting ing some of the unique chemical compounds found in the lichen’s because these lichens represent a fungal tissue. There is still much more to learn about these yeasts and symbiotic relationship between three their role in the lichen relationship. kingdoms of life: fungi from King- dom Fungi, green algae from King- dom Protista, and cyanobacteria on their ability to exist on bare rock Unlike plants, lichens have no from Kingdom Eubacteria (formerly in the baking desert or survive expo- barriers to protect themselves from Kingdom Monera). Additionally, sure to the extremes of outer space. extreme conditions. They have no like many other organisms, lichens However, they are actually quite sen- waxy cuticle or stomata to prevent have numerous species of bacteria sitive to environmental conditions. water from leaving or pollutants that live on and in them. Lichens are passive organisms and from entering the lichen. They don’t The lichen relationship is an ex- they have few mechanisms to miti- have roots either. There is no active ample of a mutualistic symbiotic re- gate changes in their surroundings. uptake of water or nutrients in li- lationship—the partners benefit from living together. The fungal partners provide general protection for the photo-partner. This includes a structural ‘home’ within which the photo- partner lives. Chemical compounds found in fungal tissue are unpalat- able or even toxic to some critters, thereby protecting the photo-partner from herbivory. Other chemical compounds in fungal tissue function as ultraviolet filters which protect the alga or cyanobacteria from the sun’s harmful rays. In return, the photo-partner feeds the fungi by producing sugars through photosynthesis. In addition to producing sugars, lichens that contain cyanobacteria (collectively called cyanolichens) also transform atmospheric nitrogen into a form that the fungi can use. Living together as a lichen, the partners are able to colonize a much wider range of habitats than if they were on their own. Lichens are valuable as biological indicators because they are respon- Lichens can be categorized into three general growth forms, similar to how we group sive to environmental change. Li- plants as herbs, shrubs, and trees. The lichen growth forms are: 1) foliose= leafy, with a chens give the general impression of distinct upper and lower surface; 2) fruticose= shrubby or hair-like; and 3) crustose= thin being tenacious little organisms based crust tightly attached to its substrate.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 3

V45_2_book.pmd 3 6/9/17, 6:52 AM ing the false impression that they are tough organisms. The composite nature of lichens is a key factor in their sensitivity to environmental change. The symbiotic relationship between the fungi and photo-partner is relatively fragile. When the lichen is stressed—by pollution or climate extremes—the balance between the partners is up- set. Usually, the stressor affects the physiology of the photo-partner, thus disrupting photosynthesis. The result is that the relationship falls apart and the lichen dies.

AIR QUALITY Lichens were first used as indicators of air quality in the early 1800s when the onset of the Industrial Revolution brought a marked increase in coal burning. People across Europe documented the disappear- ance of lichens in urban areas as pollution levels rose. Lichens can be used to indicate a variety of different pollutants including semi-vola- tile organic compounds, radionu- clides, metals, ozone, and fluorine, as well as nitrogen- and sulfur-containing compounds. Lichens are being used to monitor nitrogen-based pollutants across the country as part of a US Forest Service program that tracks forest health (Jovan 2008). Two such pollutants are nitrogen oxide and am- monia, which result from fossil fuel combustion and agricultural activity. There is an increasing interest in TOP: Cyanolichens, such as the lungwort lichens (Lobaria species), are sensitive to pollution using lichens to monitor nitrogen and their presence in an ecosystem indicates clean air. • BOTTOM: Sunburst lichens, orange foliose lichens in the genera Xanthoria and Xanthomendoza, are broadly considered nitrogen- pollution because of the growing loving species. concern over the ecological effects of nitrogen deposition. These effects chens. They are at the mercy of the Lichens are biologically active include the following: reduced atmosphere for providing moisture, only when they are wet—that’s when drinking water quality, eutrophica- along with dissolved nutrients, the photo-partner is photosynthe- tion of estuaries, nitrate-induced which are passively absorbed across sizing and making sugars, and the toxic effects on freshwater biota, the entire surface of the lichen body. fungi are growing or making struc- changes in plant community com- The lichen has no way of discrimi- tural repairs. When the lichen dries position, disruption in nutrient cy- nating between nutrients and pol- out, it goes into a state of suspended cling, and increased emissions from lutants. So, pollutants dissolved in animation and no metabolic activity soil of nitrogenous greenhouse gas- rain water are also absorbed by the occurs. In this dry state, lichens can ses (Fenn et al. 1998). lichen. withstand extreme conditions, giv- An easy, low-cost method for

4 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 4 6/9/17, 6:52 AM Fog lichens (Niebla homalea) form small, shrubby tufts, typically on coastal rocks. Their shrubby growth form intercepts wind-blown fog, enabling these lichens to become hydrated and begin photosynthesizing without a rainfall event.

monitoring air quality is to assess 3-8 kg N ha-1 yr-1 for mixed conifer of our changing climate (Jovan 2008, lichen community composition forests, chaparral, and oak wood- Geiser and Neitlich 2007, Van Herk (which species are present and their lands (Fenn et al. 2010). Lichens 2002). Lichens, like plants, respond abundance). Some lichens are very are among the most sensitive organ- predictably along climate gradients. sensitive to nitrogen and begin to isms to nitrogen pollution, so keep- They sort themselves into geo- die-off when nitrogen levels rise. ing deposition amounts below the graphic zones that share similar tem- Other lichens are nitrogen-loving critical load is expected to protect perature and moisture conditions. and flourish in high nitrogen envi- all components of these ecosystems Temperature is an important fac- ronments. As nitrogen deposition from adverse effects of nitrogen. tor in explaining lichen distribution increases, there is a distinct shift Nitrogen deposition can be mea- because it affects the rate of photo- in lichen community composition: sured using mechanical air air qual- synthesis and basic metabolic pro- nitrogen-sensitive species drop out ity sensors. However, the benefit of cesses. Going from the California of the picture and nitrogen-loving using biological indicators is that coast to the inland mountains you species move in. By monitoring the they tell you where pollutants are cross several temperature zones, proportion of nitrogen-sensitive spe- affecting the ecosystem—locations each characterized by a unique as- cies to nitrogen-loving species in an where the lichen community com- semblage of lichens. The relatively area, we can estimate the amount of position is changing. Additionally, moderate temperature at the coast nitrogen deposition. lichen monitoring can be conducted supports a lichen community that is Lichen-based estimates for nitro- throughout the landscape; whereas, distinctly different from those found gen deposition are backed by a quan- mechanical sensors are restricted to in warmer areas like the Central Val- titative benchmark. Research was areas with good access for installing ley or cooler areas like the Sierra done within California’s major for- and maintaining the equipment. Nevada Mountains. est and shrub vegetation types to As the climate warms, these tem- determine the threshold for nitro- CLIMATE perature zones will change in size gen deposition, above which the li- and location—so, too, will the dis- chen community begins to shift to- Within the last two decades a tribution of lichen communities. Li- wards nitrogen-loving species. This growing body of literature shows chens restricted to cooler forests, benchmark is known as the critical that lichens are sensitive to climatic currently found at mid to high el- load. The critical load ranged from conditions and are good indicators evations, will likely experience a

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 5

V45_2_book.pmd 5 6/9/17, 6:52 AM Methuselah’s beard, Usnea longissima, indicator of old growth forest habitat. Photograph by Richard Droker.

shrinking range. Provided the spe- requires frequent periods of wet- regimes, lichen physiology is di- cies are not dispersal-limited, they ting while others can stand regular rectly affected and we expect to see are expected to move up in eleva- and long episodes of dehydration. corresponding shifts in the lichen tion or migrate northward in search As climate change alters moisture community composition. of more suitable One particular growing conditions. group of lichens Lichens are also stands out as being closely tied to mois- intimately linked to ture regimes due to coastal climate. The their passive fog lichens, those in method for obtain- the genus Niebla (the ing water and nutri- genus name means ents. They only fog in Spanish), are colonize habitats found only in mari- with atmospheric time regions. Aside conditions that meet from one species that their hydration stretches up into needs. The physiol- Green stubble lichen (Calicium viride), indicator of old growth forest habitat. Alaska, North Amer- ogy of some species Photograph by Eric Peterson. ica’s fog lichens are

6 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 6 6/9/17, 6:52 AM concentrated along the California give a twig the appearance of a five excess in North American ecosys- coast from Sonoma County south o’clock shadow. One species of pin tems: Predisposing factors, ecosys- into Baja California. As the name lichen, C. adspersum, is listed as rare tem response, and management indicates, these lichens are restricted in California (CRPR 2B.2). While strategies. Ecological Applications 8: to a narrow section of shoreline habitat specificity is what makes 706-733. within the fog zone—an area that these lichens good indicators of old Fenn, M.E., E.B. Allen, S.B. Weiss, S. Jovan, L.H. Geiser, G.S. Tonnesen, receives a significant amount of sum- growth stands, it also means that R.F. Johnson, L.E. Rao, B.S. Gimeno, mer fog. During the summer when their fate is inextricably linked to F. Yuan, T. Meixner, and A. Bytnero- there is little to no rain, fog lichens the survival of that habitat. wicz. 2010. Nitrogen critical loads depend on regular moisture input Another indicator of old growth and management alternatives for N- from fog. forest is Methuselah’s beard, Usnea impacted ecosystems in California. The fate of fog lichens is uncer- longissima. This species is also rare Journal of Environmental Management tain in the face of climate change. in California (CRPR 4.2). In con- 91:2404–2423. California’s climate models don’t trast to the tiny pin lichens, U. Geiser, L.H., and P. Neitlich. 2007. Air agree about predictions for coastal longissima is California’s largest li- pollution and climate gradients in fog. If the frequency or duration of chen. It hangs from old growth western Oregon and Washington in- fog decreases, it will likely lead to a conifers and hardwoods with gar- dicated by epiphytic macrolichens. reduction of fog lichens. land-like strands typically reaching Environmental Pollution. 145: 203– more than a meter in length. 218. Opening your eyes to lichens can Jovan, S. 2008. Lichen bioindication of OLD GROWTH FORESTS biodiversity, air quality, and climate: be personally fulfilling. Also, the baseline results from monitoring in Old growth forests and wood- stories the lichen community has to Washington, Oregon, and California. lands are increasingly rare. In order tell can inform environmental policy General Technical Report PNW- to preserve biological diversity, con- and ecosystem management. But the GTR-737. US Department of Agri- servationists are interested in pro- first step toward seeing and learn- culture Forest Service, Pacific North- tecting as many remaining old ing from lichens is to increase li- west Research Station. growth stands as possible. To do chen awareness. The California Li- Rose, F. 1976. Lichenological indica- that, they have developed a number chen Society’s effort to designate tors of age and environmental con- of tools to rank stands in terms of lace lichen (Ramalina menziesii) as tinuity in woodlands. In Brown, importance. One tool uses lichens the California state lichen was a D.H., D.L. Hawksworth, and R.H. as indicators of stand age and conti- significant move in that direction. Bailey, (eds), Lichenology: Progress nuity (Rose 1976). Mature forests On January 1, 2016 California be- and Problems: 278–307. London, etc.: develop an architecture not found came the first state to list a lichen Academic Press. Selva, S.B. 2003. Using calicioid lichens in younger stands. Most importantly, as a state symbol. With this impor- and fungi to assess ecological conti- they include a range of tree sizes tant public acknowledgement of an nuity in the Acadian Forest Eco- and ages. This structure provides an often overlooked group of organ- region of the Canadian Maritimes. array of unique microhabitats and isms comes a tremendous potential The Forestry Chronicle, 79(3): 550- microclimates that in turn support to reach California’s school-age kids 558. unique groups of lichens. The over- as they learn about the state’s sym- Spribille, T., V. Tuovinen, P. Resl, D. all lichen diversity, in combination bols. Lichens now have a place Vanderpook, H. Wolinski, M.C. with the frequency of old growth alongside the California poppy, red- Aime, K. Schneider, E. Stabentheiner, associated species, is used to evalu- wood, and grizzly bear. Today, more M. Toome-Heller, G. Thor, H. ate stand quality. people than ever are seeing lichens. Mayrhofer, H. Johannesson, and J.P. A group of very tiny lichens called Naturalists young and old, scien- McCutcheon. 2016. Basidiomycete pin lichens, those in the genus tists, and policy makers are begin- yeasts in the cortex of ascomycete macrolichens. . Published Calicium, are broadly known as old ning to understand the impor- Science online July 21, 2016. doi: 10.1126/ growth indicators (Selva 2003). Even tant role lichens play in the envi- science.aaf8287. lichenologists have a hard time see- ronment. Van Herk, C.M., A. Aptroot, and ing these lichens because you need H.F. Van Dobben. 2002. Long-term to have your nose within two inches REFERENCES monitoring in the Netherlands sug- of them before you can see their gests that lichens respond to global characteristic 1-2 mm tall pin-like Fenn, M.E., M.A. Poth, J.D. Aber, J.S. warming. The Lichenologist, 34, pp stalks. Pin lichens, also called stubble Baron, B.T. Bormann, D.W. Johnson, 141-154. lichens, grow on bark or dead wood A.D. Lemly, S.G. McNulty, D.F. Ryan, and a dense patch of the lichen can and R. Stottlemyer. 1998. Nitrogen Shelly Benson, [email protected]

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 7

V45_2_book.pmd 7 6/9/17, 6:52 AM LICHEN CONSERVATION IS COMPLICATED by Rikke Reese Næsborg

he conservation aim for rare, threatened, or endangered (hereafter, “rare”) organisms is to maintain populations, Tso that they are able to sustain themselves. This can be a difficult task even for well-known populations of large, showy-flowered vascular plants, but the task is far more daunt- ing for inconspicuous, poorly understood taxa, such as lichens. Con- servationists generally strive to include as many different organisms as possible when designating land for conservation. However, they face special challenges with lichens because they are composite organisms that often occupy different habitats than other rare taxa.

THREATS Lichens in general are sensitive Peltigera gowardii grows entirely submerged in fresh water and is one of the few aquatic to changes in their environment. This lichens. It is associated with cyanobacteria that give it a translucent, dark green or brownish color under water. Peltigera gowardii grows in pollution-free, cold-water mountain creeks sensitivity can be attributed to three with little or no sediment or disturbance. It is endemic to North America and threatened properties of lichens. (1) They do by hydrological changes, recreational activities, erosion, and logging (Poulsen and Carlberg not have roots to absorb water and 2007). Photograph by Rikke Reese Næsborg.

WHY CARE ABOUT LICHENS? Lichens serve many functions in the ecosystem. • Some lichens are considered pioneer species capable of colonizing and breaking down bare rock. Lichens trap dust and silt as they grow, and when they die organic material is added to the minerals of the rock (Brodo et al. 2001). These processes nucleate soil formation that fosters growth of other organisms such as vascular plants. • Lichens contribute nutrients to the ecosystem by adding organic material to the soil when they decompose. Those with a cyanobacterial partner can bind nitrogen from the air, so they are important components of the nitrogen cycle. As much as 50% of the total nitrogen input in some old-growth forests has been attributed to lichens (Brodo et al. 2001). • Lichens serve as a food source for a wide variety of animals, including muskoxen, deer, voles, flying squirrels, slugs, and mites. Up to 90% of a reindeer’s winter diet may consist of lichens (Brodo et al. 2001), and many birds use lichens as a nesting material. Invertebrates such as water bears live in the intricate microtopography of lichens. • Many lichens contain unique chemical compounds that serve functions such as reflecting excess sunlight, aiding in gas exchange, or deterring herbivory. Some of these chemical compounds even have properties that benefit humans, such as antibacterial, antibiotic, cytotoxic, antiviral, and antitumor functions. Some lichen compounds are promising novel sources of natural non-genotoxic anticancer drugs (Molnár and Farkas 2010).

8 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 8 6/9/17, 6:52 AM nutrients from their substrates, nor do they have a waxy cuticle to con- HOW DO CALIFORNIA LICHENS OFFICIALLY ATTAIN trol hydration. Instead, lichens pas- “RARITY” STATUS? sively absorb water and nutrients from their environment, which con- alifornia Native Plant Society (CNPS) partners with the Califor- sequently also leads to absorption of Cnia Lichen Society (CALS) to promote conservation of rare, atmospheric pollutants that happen threatened, and endangered lichens. Information about rare lichens to land on their surfaces (McCune is collected by CALS Conservation Committee, submitted to the and Geiser 2009). (2) Each lichen is California Natural Diversity Database, and subsequently included in a composite organism, consisting one CNPS’s Inventory of Rare and Endangered Plants. Each lichen taxon or more fungi (Spribille et al. 2016) proposed as a Lichen of Conservation Concern undergoes a rigorous in a symbiotic relationship with sponsorship process by CALS to understand taxonomy, biogeogra- green algae and/or cyanobacteria. phy, and threats before status is recommended. The sponsorship The lichen symbiosis is delicate and includes a 90-day technical review, followed by publication of a therefore vulnerable to dissociation. comprehensive status review for comments by both experts and Changes in environmental condi- amateurs. Because lichen biogeography is poorly understood, a one- tions, especially increased pollution, year waiting period is required before final decisions about ranking can alter the symbiotic balance between the fungi and the algae/ and listing are made (Sims et al. 2014). cyanobacteria, resulting in a break- As of March 2017, the following 13 lichens were registered in the down of the relationship (McCune CNPS inventory: Calicium adspersum, Cladonia firma, Graphis and Geiser 2009). (Read more about saxorum, Hypogymnia schizidiata, Mobergia calculiformis, Peltigera air pollution and lichens in the ar- gowardii, Ramalina thrausta, Solorina spongiosa, Sulcaria isidiifera, ticle “Seeing Lichens” in this issue). Sulcaria spiralifera, Texosporium sancti-jacobi, Thamnolia vermicularis, (3) Many lichens are slow to colo- and Usnea longissima. nize new habitats due to long generation times and limited dispersal distances, rendering them sensitive Because of these properties, li- anthropogenic influences, such as to landscape-level disturbances chen-rich habitats worldwide are habitat degradation or loss, habitat (Sillett et al. 2000). negatively affected or destroyed by fragmentation (Bergamini et al., 2005), climate change (Ellis et al., 2007), and air pollution (Jovan 2008). Of these, habitat loss, especially of virgin forest (Wolseley 1995), is probably the most devastating threat to lichens because local populations are removed. Since most forests reach economic maturity long before they reach ecological maturity, naturally heteroge- neous forests in many parts of the world are converted to monospe- cific, even-aged stands with short rotation periods to maximize economic profits (Thor 1999). The effect on both species richness and community composition of lichens Hypogymnia schizidiata was described as a new species in 2002, and it is considered rare is dramatic and devastating. Lichen throughout its limited range. Like other tube lichens, it has hollow branches, but the floras of natural forests are unique; surface of these branches characteristically disintegrates into flakes, giving it a “schizidiate” similar communities are not found appearance. The species grows on bark and exposed wood of both conifers and hardwoods in secondary or fragmented forests in woodlands, forests, and chaparral (McCune 2006). Until recently, the known global (Bergamini et al., 2005). These distribution was limited to four islands: Cedros and Guadalupe Islands in Mexico, and Santa Cruz and Santa Rosa Islands in California. In 2015 it was discovered on the San unique lichen communities not only Francisco Peninsula. Destruction of habitat by fire may be the greatest threat in California. depend on the habitat and moisture Photograph by Rikke Reese Næsborg. regimes found in virgin forests, but

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 9

V45_2_book.pmd 9 6/9/17, 6:52 AM they also rely on long-term ecologi- spicuous organisms (Scheidegger cal continuity for dispersal and es- and Werth 2009). tablishment. Compared to more well-known Structural changes in the agricul- organisms such as vascular plants, tural landscape and urbanization mammals, or birds, a large propor- also pose serious threats to lichens. tion of lichen taxa have cosmopoli- Large-scale modern agricultural tan distributions. Most of these wide- practices eliminate minor and mar- spread lichens are unevenly distrib- ginal habitats such as remnant trees, uted throughout their range, but they stone walls, alleys, and old barns frequently receive low conservation that serve as refugia for lichens in a priority even in parts of the world changing environment (Thor 1999). where they are truly rare. Genetic Furthermore, nitrogen compounds research into a given species may generated by agricultural activities reveal that there are actually two or and by combustion of fossil fuels more geographically distinct taxa, are known to have harmful effects such that lack of protection could on lichens (McCune and Geiser reduce or even extirpate genetic di- 2009). versity (Scheidegger and Goward Ramalina thrausta resembles a tiny, pen- 2002). Some lichens are rare through- dent shrub with filamentous branches that out their widespread range. For ex- SPECIFIC PROBLEMS WITH are curled over at the tips. these curly tips have earned it the poetic common name ample, species of Lobaria, Sticta, Ne- CONSERVATION OF “angel’s hair.” The species is mainly found phroma, Pannaria, and Calicium are LICHENS on dead twigs of both conifers and hard- associated with boreal and temper- woods in moist, cool forests of the boreal ate forests that are open and ancient. Since each lichen is a composite zone, and it is considered uncommon Previously, these forests were kept throughout its global range. It is mainly organism, lichen conservation must threatened by air pollution and timber open by populations of large, graz- deal with at least two very different harvest (Carlberg 2006). Photograph by ing herbivores, but a combination of species. Even though the fungus is Tom Carlberg. declining or disappearing large graz- typically the object for conserva- ers as well as logging of ancient for- tion efforts (Scheidegger and degger et al. 1995; Sillett et al. 2000), ests has rendered these habitats in- Goward 2002), the delicate nature but this conservation approach will creasingly scarce. In many areas, the of the symbiosis creates challenges be inherently more challenging above mentioned lichens are now unique to lichens. Developing con- for sexually reproducing lichens restricted to manmade parks, wood servation strategies for lichens is ad- because their reproductive units pastures, and hunting reserves that ditionally complicated by the fact (spores) contain only the asco- have remained unchanged for many that the distributions and ecologies mycete fungus, and also for crus- years (Wolseley 1995). Some lichen remain unknown for most lichen tose species because of their small genera are narrow endemics, and taxa. Moreover, many lichens, par- sizes. Conservation must instead be there are also many local, endemic ticularly inconspicuous crustose based on strategies that protect suit- species within widespread genera. species, await scientific description able habitats for the long-term, but Endemic species may either repre- or have yet to be discovered alto- the lack of knowledge about lichen sent emergence of “new” taxa via gether. This is probably especially ecology means these decisions of- speciation or reflect relict survivors true for rare taxa. ten rely on guesswork. Management after extensive range reduction (Gal- While it is possible to cultivate decisions are further complicated loway 1997). Regardless of origin, and breed many other species in by the fact that many lichens have rare endemics are often considered captivity (e.g., Hanson and Nelson specific habitat requirements that high conservation priorities—that is, 2015), the delicate symbiosis of li- are not shared by other organisms if we know enough about their dis- chens is poorly understood and no- in the same community, so special tributions and ecologies to offer ef- toriously difficult to duplicate in lichen-specific management consid- fective protection measures. captivity (Honegger 1996) which erations may be needed. As a result, prohibits our ability to artificially lichen preferences are often not CONSERVATION augment lichen populations. Trans- taken into account when planning STRATEGIES plantation of symbiotic propagules reserves, and an overall ecosystem and lichen fragments can be suc- approach of conservation often fails In order to maintain viable popu- cessful for some species (Schei- to successfully protect these incon- lation sizes over time it is vital that

10 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 10 6/9/17, 6:52 AM current large populations of rare li- REFERENCES view. Zeitschrift für Naturforschung chen species receive strict protec- 65: 157–173. tion. Such large populations serve Bergamini, A., C. Scheidegger, S. Stofer, Poulsen, B. and T. Carlberg. 2007. P. Carvalho, S. Davey, M. Dietrich, as propagule sources for expanding Peltigera hydrothyria, Sponsorship F. Dubs, E. Farkas, U. Groner, K. the ranges and therefore enhancing for the CALS Conservation Commit- Karkkainen, C. Keller, L. Lokos, S. tee. Bulletin of the California Lichen the long-term survivability of rare Lommi, C. Maguas, R. Mitchell, Society 14:15–18. species. Of course, the success of P. Pinho, V.J. Rico, G. Aragon, A.M. Scheidegger, C., B. Frey, and S. Zoller. these conservation efforts is only Truscott, P. Wolseley, and A. Watt. 1995. Transplantation of symbiotic possible if adjacent or nearby future 2005. Performance of macrolichens propagules and thallus fragments: habitats for range expansion are ap- and lichen genera as indicators of Methods for the conservation of propriately managed, such that suit- lichen species richness and compo- threatened epiphytic lichen popula- able habitats (e.g., old trees for epi- sition. Conservation Biology 19: tions. Mitteilungen der Eidgen- phytes or open soil patches for soil- 1051–1062. ssischen Forschungsanstalt für Wald, dwelling species) will continue to Brodo, I.M., S.D. Sharnoff, and S. Schnee und Landschaft 70:41–62. be available for colonization within Sharnoff. 2001. Lichens of North Scheidegger, C. and T. Goward. 2002. America. Yale University Press, New the landscape (Scheidegger and Monitoring lichens for conservation: Haven and London. red lists and conservation action Werth 2009). Often, it is necessary Carlberg, T. 2006. Ramalina thrausta. plans. In: Lichen Monitoring—Moni- to considerably reduce or altogether Conservation Committee Sponsor- toring Lichens. Ed. P.L. Nimis, C. halt harvest of forest resources since ship. Bulletin of the California Lichen Scheidegger, P. Wolseley. Kluwer many lichens are extremely sensi- Society 13: 17. Academic Publishers. 163–181. tive to environmental changes Ellis, C.J., B.J. Coppins, T.P. Dawson, Scheidegger, C. and Werth, S. 2009. (Jovan 2008). and M.R.D. Seaward. 2007. Response Conservation strategies for lichens: A successful conservation strat- of British lichens to climate change insights from population biology. egy typically combines habitat pro- scenarios: trends and uncertainties Fungal Biology Reviews 23:55–66. tection with efforts to maintain or in the projected impact for contrast- Sillett, S.C., B. McCune, J.E. Peck, T.R. improve the size, distribution, and ing biogeographic groups. Biological Rambo, and A. Ruchty. 2000. Dis- Conservation 140:217–235. genetic structure of lichen popula- persal limitations of epiphytic li- Galloway, D.J. 1997. Lichen Biogeog- chens result in species dependent on tions. This does not necessarily mean raphy. In: Lichen Biology. Ed. T.H. old-growth forests. Ecological Appli- that habitats, forest stands, or even Nash III. Cambridge University cations 10:789–799. entire landscapes must be com- Press, Cambridge. Sims, A.E., R. Bittman, E. Peterson, and pletely off limits, as long as the li- Hanson, J. and J.K. Nelson. 2015. Rare T. Carlberg. 2014. Lichens Added to chen populations can retain viable plant management success stories in the CNPS Inventory from CALS Li- effective population sizes and popu- California. Fremontia 43: 14–19. chens of Conservation Concern. lations in different areas are able to Honegger, R. Morphogenesis. In: Li- Bulletin of the California Lichen Soci- interbreed to maintain genetic di- chen Biology. Ed. T.H. Nash III. Cam- ety 21:5–12. versity (Scheidegger and Werth bridge University Press, Cambridge. Spribille, T., V. Tuovinen, P. Resl, D. 2009). Large populations with high Jovan, S. 2008. Lichen Bioindication of Vanderpool, H. Wolinski, M.C. Biodiversity, Air Quality, and Cli- Aime, K. Schneider, E. Stabentheiner, genetic variability are expected to mate: Baseline Results From Moni- M. Toome-Heller, G. Thor, H. have a higher probability of survival toring in Washington, Oregon, and Mayrhofer, H. Johannesson, and J.P. as well as a higher evolutionary po- California. General Technical Report McCutcheon. 2016. Basidiomycete tential to cope with future environ- PNW-GTR-737. US Department of yeasts in the cortex of ascomycete mental changes. Agriculture, Forest Service, Pacific macrolichens. Science 353:488–492. Despite the reality that lichen Northwest Research Station, Port- Thor, G. 1999. Red-listed lichens. In: conservation is faced with many land, OR. Nordic Lichen Flora Vol. 1. Ed. T. Ahti, challenges such as limited knowl- McCune, B. 2006. Hypogymnia schizi- P.M. Jørgensen, H. Kristinsson, R. edge about ecology and distribu- diata, Sponsorship for the CALS Moberg, U. Søchting, and G. Thor. 9– tion as well as habitat loss and Conservation Committee. Bulletin Bohuslan ‘5, Uddevalla, Sweden. 11. fragmentation, progress is being of the California Lichen Society 13: 42–44. Wolseley, P.A. 1995. A global perspec- made here in California (see side- McCune, B. and L. Geiser. 2009. tive on the status of lichens and their bar on page 9). On January 1, Macrolichens of the Pacific Northwest. conservation. Mitteilungen der Eidgen- 2016 California was the first of the 2nd edition. Oregon State University ssischen Forschungsanstalt für Wald, United States to designate a state Press, Corvallis, OR. Schnee und Landschaft 70:11–27. lichen (lace lichen, Ramalina men- Molnár, K. and E. Farkas. 2010. Cur- ziesii) to promote awareness about rent results on biological activities Rikke Reese Næsborg, rikke.reesenaesborg lichens. of lichen secondary metabolites: a re- @gmail.com

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 11

V45_2_book.pmd 11 6/9/17, 6:52 AM Bryophytes in all their glory, growing in the North Coast redwood forest. All photos by Marie Antoine. A BIT ABOUT BRYOPHYTES by Marie Antoine

lthough there is much over- The word “bryophyte” translates relatively soon after plants colonized lap in the ecology of li- to “moss plant.” Mosses are indeed land. For the past 400 million years chens and bryophytes, bryophytes, but liverworts and horn- or so, bryophytes have been adapt- they are fundamentally worts are also bryophytes. Techni- ing and evolving to exploit a broad differentA groups of organisms. A li- cally speaking, bryophytes are an range of habitats where they can avoid chen is a symbiotic system where a informal group composed of three competition with tracheophytes. fungus farms a population of green separate phyla: Bryophyta (mosses), While mosses, liverworts, and algal and/or cyanobacterial cells. In Marchantiophyta (liverworts), and hornworts are only distantly related other words, lichens represent a Anthocerophyta (hornworts). These to each other, there are certain fea- highly successful fungal nutritional are ancient land plant lineages, and tures that unite them. Bryophytes mode. Bryophytes, on the other their divergence from tracheophytes are often referred to as “non-vascu- hand, are plants. (lignified vascular plants) occurred lar plants” because they lack ligni-

12 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 12 6/9/17, 6:52 AM fied vascular tissue. Lignins impart not differentiated into leaves and structural support and rigidity in stems. Thalli may be strap-shaped the conducting tissue of tracheo- or form a rosette. If the plant is phytes, which include all other land reproductive, the appearance of the plants except bryophytes. The lack sporophyte is also diagnostic. of lignin means bryophytes can only Mosses are the most familiar be relatively small plants. Bryophytes group of bryophytes, and with over anchor to their substrates with hair- 13,000 species they are also the most like rhizoids, instead of roots. All diverse. All mosses have leafy game- bryophytes lack seeds; they repro- tophytes and their leaves are spi- duce sexually via spores and asexu- rally arranged upon the stem. ally via fragmentation or produc- Many mosses have a flattened, tion of tiny clonal propagules called feather-like appearance, but exami- gemmae. Furthermore, all bryo- nation with a hand lens will show phytes require liquid water for fer- the leaves are indeed spirally ar- tilization because of their swimming ranged. The moss sporophyte con- sperm. sists of a small brownish spore cap- Seeds and sperm may be familiar sule attached to the parent gameto- concepts, but several of the other phyte via an unbranched stem. The features uniting bryophytes involve sporophyte stem is wiry and du- less intuitive aspects of their life rable, and it often persists after the cycle. All land plants have the same spore capsule has shed its spores or fundamental life cycle of alternating been picked off by some hungry ani- heteromorphic multicellular genera- mal. The spore capsule itself has a tions: the haploid generation (with neat little lid that pops off when the one set of chromosomes) is the ga- spores are mature. In most mosses, metophyte that makes gametes (eggs the mouth of the spore capsule is or sperm) and the diploid genera- lined with tiny intricate teeth that tion (with two sets of chromosomes) flex with changes in humidity and is the sporophyte that makes spores. thereby promote spore dispersal. Bryophytes have a dominant game- Liverworts are the second most tophyte, and their sporophytes are diverse group of bryophytes with short-lived, unbranched, and nutri- approximately 5,200 species. Most tionally dependent on the gameto- liverworts are leafy and could be phyte. This situation is reversed in mistaken for mosses upon first im- tracheophytes, where the sporophyte pression. However, close examina- TOP: Kindbergia showing “feather moss” is dominant, relatively long-lived, tion of a leafy liverwort will reveal morphology. • BOTTOM: Thalloid (1) vs. leafy and branched, while gametophytes that, unlike mosses with their spi- (2) bryophytes. are variously reduced and retained rally arranged leaves, liverwort on the sporophyte. In other words, leaves are arranged in two or three dorsal air pores. In contrast to com- the green fuzzy moss is a gameto- distinct rows. There are two lateral plex thalloid liverworts, simple thal- phyte, but the green leafy tree is a rows of leaves, and most leafy liver- loid liverworts are only a few cell sporophyte. worts also have a third row of leaves layers thick and their fragile trans- on the ventral surface (not surpris- lucent bodies do not have differenti- ingly, these are called underleaves). ated tissues or dorsal air pores. HOW CAN YOU TELL Although more than 85% of all Whether belonging to a leafy or a VARIOUS BRYOPHYTES liverworts are leafy, the thalloid liv- thalloid liverwort, the liverwort APART? erworts are probably more familiar sporophyte is uniformly ephemeral. to most people because of their A fragile, transparent stem expands The first thing to do is determine prevalence as greenhouse or nurs- via turgor pressure to elevate the whether the gametophyte is leafy or ery weeds. Thalloid liverworts may shiny blackish spore capsule, and thalloid. If the plant is leafy, you be either complex or simple. Com- the whole structure collapses after will be able to see tiny leaves at- plex thalloid liverworts are many the spores are released. If you have tached to tiny stems. On the other cell layers thick, which allows for access to a compound microscope, hand, a thallus is a plant body that is tissue differentiation and turret-like squash-mount a liverwort spore cap-

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 13

V45_2_book.pmd 13 6/9/17, 6:52 AM phytism, where primitive precursors to tracheo- epiphyte means phytes. You will often see this claim “on the surface even in textbooks and scientific pa- of a plant.” Many pers, and this erroneous thinking lichens are epi- likely arises from the chauvinistic phytic as well. belief that “bigger is better.” Bryo- Both bryophytes phytes represent a fundamentally and lichens lack different strategy in land plant evo- roots. This lution. Bryophytes and tracheo- means they can- phytes diverged shortly after plants not tap into their first colonized land, and from the host’s sugar and beginning bryophytes pursued a dis- water supply parate strategy from tracheophytes. like parasitic The bryophyte strategy involved mistletoes do. elaboration of the gametophyte, and Epiphytic bryo- the tracheophyte strategy involved phytes and li- elaboration of the sporophyte. It is chens use their interesting to consider the ultimate rhizoids to cling role of the sporophyte (spore dis- to the outer sur- persal) vs. the role of the gameto- Polytrichum moss with mature sporophytes face of their host phyte (successful fertilization) and tracheophyte the characters that would be advan- sule and check out the beautiful he- and they do not penetrate their host’s tageous in each case. What makes lical structures that are mixed in bark or leaf cuticle. These epiphytes sense for a large plant does not make with the spores. These little spring- simply use their hosts as substrate. sense for a small plant, and there are like structures, called elaters, en- Growing upon the bark, wood, or some fascinating morphological, hance spore dispersal and are unique leaves of tracheophytes is an excel- anatomical, and physiological pro- to liverworts. lent strategy for reaching moisture, cesses underlying bryophyte success. Hornworts are the third and least nutrients, and sunlight without hav- The numbers speak to the success common group of bryophytes. They ing to compete directly with tra- of the bryophyte strategy: in terms are arguably a relictual group, with cheophytes for these resources. of diversity, they are second only to fewer than 200 species worldwide. If epiphytic bryophytes and li- the flowering plants. Their small All hornworts have a thalloid game- chens do not harm their substrates, size, intricacy of form, and unique tophyte, and the thallus is usually a then why do we see more epiphytes physiology make bryophytes per- distinctive bluish-green color be- on dead or dying branches? cause of symbiotic cyanobacteria Again, we must return to Thalloid liverworts Conocephalum (complex) and Pellia that live inside and provide a nitro- the issue of competition. As (simple). gen source to the hornwort. The a tree or branch dies, it loses “horn” part of the plant is the sporo- its leaves. With fewer tra- phyte, which is essentially just an cheophyte leaves, there is elongating spore capsule. Hornworts more light available for the have a number of characteristics that epiphytes. More light means make them unlike any other land more epiphyte growth, and plants. For example, each cell has voilà, the dead branch or only one large chloroplast, whereas tree appears unusually all other land plants have multiple cloaked in epiphytes. Epi- chloroplasts per cell. phyte growth may be also be enhanced by minerals WHAT ARE BRYOPHYTES leaching out of dying foli- NOT? age and twigs, but the epiphytes themselves are not First of all, bryophytes are not responsible for the tracheo- parasites, although you will often phyte’s decline. find bryophytes growing upon other Here is another myth to plants. This habit is called epi- dispel: bryophytes are not

14 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 14 6/9/17, 6:52 AM fectly suited to inhabit niches that ing organisms (including us) have are unavailable to tracheophytes. a water physiology called homeo- hydry, where cellular water content WHERE DO BRYOPHYTES must remain within certain bound- LIVE? aries or else the organism will die. Bryophytes have a type of water The short answer is: almost ev- physiology called poikilohydry. This erywhere where they can avoid com- word means “varied water.” When petition with tracheophytes. This water is available, bryophytes are competition may be either direct physiologically active. When water (imagine a moss trying to compete ceases to be available, they undergo for space with a redwood tree) or an orderly shutdown of cellular pro- indirect (imagine a deciduous tree cesses and remain dormant until wa- dropping a thick layer of leaves and ter returns. Lichens are also poi- smothering a thalloid liverwort). The kilohydric. The advantage of poiki- same concept applies to lichens, and lohydry is that bryophytes and li- this is one of the reasons why you chens are liberated from reliance on will often find bryophytes and li- soil water, so they can grow on sur- chens growing together. Generally, faces like rock and bark. The disad- bryophytes and lichens have the advantage is that these organisms will vantage on any substrate where tra- be dormant whenever ambient wa- cheophytes are disadvantaged. This ter is unavailable. It is important to includes hard impermeable sub- point out that poikilohydry and des- strates with poor water retention, iccation tolerance are not synony- such as rock, bark, and tracheophyte mous. All bryophytes and lichens leaf cuticles. Tracheophytes cannot are poikilohydric, but degree of des- usually exploit these substrates be- iccation tolerance ranges widely Leafy liverworts Porella and Frullania on cause they need to sink their roots among species from different habi- alder trunk. into a dependable water supply. Here tats. A bryophyte or lichen growing is where we encounter one of the on a desert rock can tolerate more chens thrive on hard surfaces, there profound physiological differences extreme drying-wetting cycles than are certain conditions that promote between bryophytes and tracheo- can a related species growing on a bryophyte dominance on softer sub- phytes. log in a humid forest. strates. The combination of deep Most tracheophytes and other liv- While many bryophytes and li- shade and high humidity is one such

BELOW LEFT: Leafy liverwort Scaparia (fertile, with flimsy sporophytes in variable stages of development) growing amidst a collection of mosses. • BELOW RIGHT: Hornworts with their “horn-like” sporophytes.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 15

V45_2_book.pmd 15 6/9/17, 6:52 AM bryophyte habitat. These soils often lack sufficient oxygen to support the roots of tracheophytes. Bryophytes, on the other hand, can simply absorb oxygen across their entire surface area from the air or water around them. One moss genus, Sphagnum, is considered an ecosystem engineer because of its ability to hold water and create conditions that exclude most other plants. This fascinating moss has unique anatomy and physiology that creates and perpetuates the vast northern peatlands (which is a topic deserving of its own article).

WHY SHOULD WE CARE ABOUT BRYOPHYTES?

Aside from their aesthetic charm, bryophytes have inherent value as contributors to biodiversity. They also play fundamentally important ecological roles, such as hydrological buffering and nutrient cycling. Imagine rain fall- ing through the crown of a tree that is cloaked in mosses and leafy liver- Isothecium (the icicle moss) growing on dead lower branches. worts. The bryophytes absorb water and slowly re- condition where bryophytes can reign supreme. You will find rela- lease it, like so many green sponges, colonize soil. Most bryophytes are tively few lichens in these dark, wet thereby buffering the canopy habi- considered “shade plants” in that niches. Prolonged hydration and low tat, which is otherwise quite desic- they can achieve positive carbon bal- light levels can be a dangerous com- cating. This, in turn, provides viable ance at very low light levels. An- bination for lichens. Most lichens habitat for other organisms. Every other bryophyte-favorable substrate need regular cycles of wetting and bryophyte colony is like a miniature is provided by soft, wet, decaying drying to maintain the balance of forest with niche diversity and food wood on the forest floor. With ad- the symbiosis, lest the fungus re- webs of its own. Tiny animals are equate light tracheophytes will colo- spire the system to death. Many given shelter and protection from nize this habitat, as in the “nurse bryophytes, on the other hand, will predators within the matrix of the log” phenomenon where young thrive under conditions of near- bryophyte forest. Larger animals, trees establish on fallen logs. In the constant hydration. particularly birds, use bryophytes as deep shade, however, bryophytes Waterlogged soils can be prime nesting material. Moss spore cap-

16 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 16 6/9/17, 6:53 AM sules are nutritious snacks for in- headlamp and sects, birds, and small mammals. a hand lens to explore the intricate colonies HOW ARE BRYOPHYTES of leafy liver- FARING IN THE worts on old ANTHROPOCENE? wet redwood logs. Stroll (oh- A major threat to bryophyte (and so-carefully) lichen) abundance and diversity along a stream is habitat destruction, whether bank and ad- through clear-cut logging, urban de- mire the thal- velopment, or collection for the hor- loid liverworts ticultural industry. Another impor- growing on tant vulnerability is air pollution. silty mineral Recall the poikilohydric nature of soil. Finally, search for bryophytes and lichens. All water hornworts on clay cut- and nutritional needs must be met banks or old gravel road- through direct deposition from the beds. The more you look atmosphere. While this is a handy for these enchanting little strategy for making a living literally plants, the more you will out of thin air, it also means bryo- find the “green blur” com- phytes and lichens are unable to ing into delightful and un- exclude harmful components in expected focus. their environment. Lichens are clas- sic indicators of air pollution, as certain species will disappear from FURTHER READING polluted areas and be replaced by ON GENERAL pollution-tolerant species. This is BRYOLOGY also true for bryophytes. You might recall the recent news from Port- Goffinet, B. and J. Shaw land, Oregon where street tree (eds.) 2008. Bryophyte mosses revealed a previously un- Biology. Cambridge Uni- known air pollution problem (i.e., versity Press, U.K. heavy metal deposition). Climate Kimmerer, R.W. 2003. Gath- TOP: Sphagnum, the ecosystem engineer. • BOTTOM: change is another likely threat to ering Moss: A Natural Hummingbird nest with woven moss. bryophytes, although it remains to and Cultural History of be seen how that will unfold. Mosses. Oregon State University Mosses. Micro-Optics Press, New Press, Corvallis, OR, U.S.A. Zealand. Norris, D.H. and J.R. Shevock. 2004a. WHERE MIGHT ONE LOOK Contributions toward a bryoflora FOR BRYOPHYTES? FURTHER READING FOR of California I. A specimen-based BRYOPHYTE catalogue of mosses. Madroño 51: No doubt after reading this 1-131. Fremontia issue you will be out and IDENTIFICATION Norris, D.H. and J.R. Shevock. 2004b. about looking for lichens. As you Doyle, W.T. and R.E. Stotler. 2006. Contributions toward a bryoflora of examine epiphytic lichens growing California II. A key to the mosses. on shrubs and trees, pay attention Contributions toward a bryoflora of California III. Keys and annotated Madroño 51: 133-269. to the mosses and leafy liverworts species catalogue for liverworts and that will likely be growing inter- hornworts. Madroño 53: 89-197. ONLINE mixed. As you examine rock-dwell- Malcolm, B. and N. Malcolm. 2006. ing lichens, look in cracks and fis- Mosses and Other Bryophytes—An Il- Bryophyte Chapter, California Native sures for moss colonies. Venture into lustrated Glossary, 2nd Edition. Mi- Plant Society: bryophyte.cnps.org the shady nooks and crannies of your cro-Optics Press, New Zealand. local forests and find luscious mosses Malcolm, B., N. Malcolm, J. Shevock, Marie Antoine, marie.antoine@humboldt. carpeting the forest floor. Use a and D. Norris. 2009. California edu

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 17

V45_2_book.pmd 17 6/9/17, 6:53 AM A field biologist samples a stand of tufted poppy (Eschscholzia caespitosa) in the Inner Coast Ranges of Napa County with a backdrop of repeating floristic patterns (labeled and delineated). These patterns are guided by effects of soil, exposure, climate and fire. Repeat mapping takes advantage of definitions derived from vegetation sampling to accurately depict changes over time. TIMES THEY ARE A CHANGING: VEGETATION MAPPING AND RE-MAPPING EFFORTS ACROSS CALIFORNIA by Todd Keeler-Wolf, Rachelle Boul, Jennife Buck-Diaz, and John Menke

vegetation map is a graphic of a mouse, you can display not only grasslands from highly diverse representation of the pat- the floristic variety of a given area’s patches of native wildflowers con- terns of vegetation such vegetation, but also its structural taining rare endemic plants. This as woodlands, chaparral, attributes (such as height of the liv- information helps people explore Aand grassland arrayed across a land- ing vegetation), and even habitat and explain a wide variety of bio- scape. Though vegetation maps have quality for specific animals and logical information for a mapped been produced for over 100 years, plants. For example, a GIS map of area. It is no wonder that today’s those created today far surpass a Sonoma County vegetation high- vegetation maps have become an simple two-dimensional map. Geo- lights the tallest forests of large- indispensable tool for conservation graphic Information System (GIS) diameter coast redwoods which can planning and natural resource as- technology enables maps to repre- host nest sites for the endangered sessment. sent a variety of “attributes” in addi- marbled murrelet, and can also dif- The process of repeat vegetation tion to vegetation type. At the click ferentiate largely non-native annual mapping synthesizes and summa-

18 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 18 6/9/17, 6:53 AM rizes changes for all vegetation cov- nent viewpoints such as hills and tions of historic controls on the ex- ered by the map. This confers a ridges by field mappers. This tech- tent and distribution of vegetation. unique status to vegetation mapping nique introduces a number of prob- For example, a 2010 map in the as the single best “landscape-level lems for accurate depiction of land- vicinity of the City of Oakland de- monitoring” tool amongst the arse- cover patterns. By necessity, the picts how vegetation might have ap- nal of biological monitoring meth- maps were drawn at a broad scale peared 300 years ago prior to its ods. For proper use, however, re- (less detail) to encompass the inher- first sighting by European explor- peat mapping must be carefully done ent uncertainties and to enable this ers. This view is based on the land- with appropriately set limitations. huge effort to be completed over a scape-level influence of Native This article explores the value of manageable time frame (Figure 1). American land management prac- repeat mapping through numerous Researchers interested in making tices, which was not well under- examples across California, while detailed comparisons between the stood when Kuchler developed his highlighting the careful steps that Wieslander map and current condi- maps (Figure 2). must be taken to guarantee the high- tions must first generalize newer est comparison value. maps to compare “apples-to-apples.” ORANGE COUNTY Some results have been relatively (20-YEAR INTERVAL) inconclusive, however, analysis of INTERPRETING VEGETATION the extensive field data collected to In more recent efforts, the Cali- INFORMATION FROM THE support these maps suggest broad fornia Native Plant Society compared PAST landscape level changes at the tree a 2012 vegetation map within Or- line in both lower and upper eleva- ange County (AIS 2015a) to an ex- The first vegetation map across tions in the mountains of Califor- isting map produced in 1992 (Jones California was spearheaded by A.E. nia, helping to confirm evidence of and Stokes 1993) to interpret differ- Wieslander for the US Forest Ser- climate change patterns (Thorne et ences between the maps and to guide vice in the 1930s. Its quantitative al. 2008, Wright et al. 2016). future change analyses. Natural nature and broad geographic scope, Vegetation conditions prior to lands within Orange County include covering >40% of the state, has made recorded history have also been con- habitats such as open grasslands, it an important resource to revisit sidered, perhaps the most famous of coastal sage scrub, chaparral, and (Kelly and Allen-Diaz 2005, Thorne these was produced by Kuchler coast live oak woodlands. The veg- et al. 2008, etc.). However, there are (1977) for California as a “potential etation in this region has undergone problems using these beautifully vegetation” map. This map synthe- fragmentation due to urban expan- produced maps as a basis of com- sized scientific information on the sion as well as multiple, widespread parison to current vegetation condi- status and distribution of plants be- fires causing change and succession. tions. They were made at a time fore European history in the state, The Fire and Resource Assess- when geodetic surveying (measure- presumably when natural processes ment Program (FRAP 2014) pub- ment of the earth) was less accurate were running their course without lishes a statewide geodatabase of fire and vegetation classification rules the “control of man.” Though use- perimeters that includes wildfire his- were less rigorous. There were no ful for conceptualizing past condi- tory, prescribed burns, and other aerial photographs for reference, and tions, these maps are open to inter- fuel modification projects. This re- they were hand-drawn from promi- pretation depending upon assump- source allows users to understand

FIGURE 1. A comparison of Wieslander Vegetation Type Map (VTM) (A), 2010 northern Sierra Nevada foothill vegetation map (B), and recent color aerial imagery for the same area between Chico and Paradise in the northern foothills, depicted at 1:18,000 (C). the more accurate and detailed 2010 map is a result of quantitative vegetation classification and detailed aerial photography as a basis for the mapping, neither of which were available for the VTM mapping 75 years ago. SOURCE: Wieslander, A.E. 1935. A vegetation type map of California. Madroño. 3:140–144; Menke, J., et al. 2011. Northern Sierra Nevada Foothills Vegetation Project: Vegetation Mapping Report; Google, DigitalGlobe, 2016.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 19

V45_2_book.pmd 19 6/9/17, 6:53 AM 20 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 20 6/9/17, 6:53 AM FIGURE 3. The western Riverside remap highlights significant change from coastal scrub dominated in 2001 to annual grasses in 2012 (this change is shown in yellow) in 2012 and the positional relationship to fire (shown in red) which occurred across an 11-year time span. SOURCE: Aerial Information Systems, Inc. 2015b. Western Riverside County vegetation mapping update. Final vegetation mapping report. Prepared for the Western Riverside County Regional Conservation Authority. Redlands, CA.

and interpret landscape-scale veg- reproduce and/or build up adequate (currently at low levels 2-10%), re- etation change correlated with fire seed reserves. In 2006, a large fire gardless of the overall vegetation across California. Variation in the swept over the northern-most popu- type. Supplementary attributes for frequency, intensity, and scale of fire lation of Tecate cypress, damaging each polygon allow users to identify significantly alters the patterns of the majority of the mature individu- and quantify more than just the ex- plant community regeneration. One als. The 2012 map delineates only isting vegetation type. important, rare vegetation type in 38 acres of Tecate cypress compared Mapping efforts represent single this region are stands of Tecate cy- to 196 acres delineated in 1992 snapshots in time with unique inter- press (Hesperocyparis forbesii Alli- (<20% of the 1992 extent). Much of pretations of the distribution of veg- ance), which grow along dry, ex- the area previously mapped as Tecate etation in the region and allow users posed hillsides and ridgetops, stream cypress is now dominated by vari- to better understand and interpret banks, and arroyos. This species has ous shrub species. However, signifi- vegetation patterns across the region. a California Rare Plant Rank of 1B.1 cant regeneration of the cypress has However, differences between maps and is threatened by trends of short been observed, and this regenera- are not always straightforward. Care- fire return intervals, which destroy tion is documented in the 2012 map ful map interpretation is needed to young trees before they are able to within the conifer density attribute differentiate actual changes in vegetation composition from differences FACING PAGE: FIGURE 2. A hypothetical map of vegetation in the area now occupied by the in classification, map scale, and reso- city of Oakland immediately prior to the European discovery of the San Francisco Bay lution of delineation. circa 1700. This map was created by studying remaining natural vegetation in the area and reviewing historic images and documents from the 1700s through early 1900s. A USGS topographic map backdrop shows the current arrangement of streets and landmarks. WESTERN RIVERSIDE Changes have occurred in large areas of salt marsh (Sarcocornia pacifica, shown as lavender, lower left), now built-up as bay-fill and including the Oakland Coliseum complex, native COUNTY (11-YEAR grasslands and herblands occupied much of the city core, while redwood (Sequoia INTERVAL) sempervirens) stands in the hills (orange color at upper right) still occupy roughly the same locations and area as they did 300 years ago. Riverside County has faced chal- SOURCE: Keeler-Wolf, T. 2010. unpublished map created for Oakland Museum of California. lenges similar to Orange County in-

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 21

V45_2_book.pmd 21 6/9/17, 6:53 AM in average annual precipitation, increases in agricultural and human population, as well as substantial wetland and channel reconfiguration. One advantage of this re-mapping effort was the use of the same mapping standards and floristic classification to ensure continuity between the two products (AIS 2005, AIS 2015b). This continuity allows a very accurate analysis of true vegetation change over time versus differences due to mapping tech- nique. Change was detectable across both floristic and structural characteristics of vegetation and can be re- ported using various scales of classification, from the alliance level up to broader types such as those defined in a regional Multi Species Habitat Conservation Plan. For example, a loss of almost 5000 acres of Brittlebush scrub (Encelia fari- nosa Alliance) be- FIGURE 4. Select areas of California have been mapped and remapped at varying intervals to assess change tween 2001 and over time. 2012 can also be ag- Source: Boul, R. 2016. gregated within the more general cat- cluding extensive wildfire and ur- This area supports coastal scrub, egory of the Coastal Sage Scrub ban expansion. The area of focus for chaparral, grasslands, riparian cor- Group which showed a loss of re-mapping included over one mil- ridors, and wetlands. Dynamics af- 13,795 acres. lion acres of western Riverside fecting vegetation change across the Another illustration of change County between the San Jacinto and 11 year interval include fire across over time includes stands of chamise Santa Ana Mountains (Figure 4). 119,000 acres, a significant decrease (Adenostoma fasciculata Alliance)

22 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 22 6/9/17, 6:53 AM that burned after the origi- coast live oak trees in the nal 2001 vegetation map. adjacent nearby grass- While these stands may re- lands. The final product main within the chamise depicts a total of over Alliance, other attributes FIGURE 5. Vigorous spread of common reed (Phragmites australis) 2,500 acres of forest and capture structural change from 1999 to 2015 in the Grizzly Island Wildife Area complex, woodlands that have been in shrub cover, e.g. from along the edge of Suisun Bay in Benicia, CA. reclassified from vegeta- 60% shrub cover to 10- SOURCE: Vegetation Classification and Mapping Program tion containing pure and (VegCAMP). In prep. 2015 vegetation map update for Suisun Marsh, 25% shrub cover. Changes Solano County, California. A report to the California Department mixed tanoak to other in shrub cover have direct of Water Resources. California Department of Fish and Wildlife, vegetation types where impacts in relation to habi- Sacramento, CA. tanoak is not a compo- tat preferences for various nent of the canopy. The wildlife species. Other noted resent the response of vegetation to final product incorporates the in- changes include post-fire increases this devastating pathogen (AIS formation gathered from each of the in levels of non-native plant spe- 2015c). three mapping efforts over the 10- cies, e.g. from category 0 (no exot- In 2004, AIS mapped nearly year period into one coherent ics observed) to category 2 (high 19,000 acres of mixed hardwoods, geospatially referenced database, exotics). By implementing quanti- redwood and Douglas-fir forests, giving biologists and other users ef- tative analyses for multiple aspects serpentine and mixed chaparral, fective decision-making tools for the of vegetative change between the maritime coastal scrub and native management of habitat loss and 2001 and 2012, informed manage- grasslands within the Mount Tamal- change, increased fire danger, and ment decisions can be made based pais watershed. This important re- other hazards resulting from stand- on floristic characteristics as well gion is located in the middle of a ing dead and downed trees. as structural attributes and the over- diverse belt of mixed broadleaf and all quality of vegetation in the re- conifer forest, paralleling the San SUISUN MARSH (7 MAPS / gion. Andreas Fault between the Bolinas 3-YEAR INTERVALS) Ridge and Ross Valley. Vegetation was mapped in stands one acre or The vegetation of Suisun Marsh MOUNT TAMALPAIS larger and assigned a vegetation type is likely the most mapped vegeta- WATERSHED (4 MAPS / as well as attributes such as cover tion in the entire state of California. 5- TO 10-YEAR INTERVALS) density of conifers and hardwoods The current methodology, adopted as well as understory shrub cover in 1999, utilizes the National Veg- The Marin Municipal Water Dis- where visible. etation Classification (USNVC) trict developed and updated a Veg- By measuring the cover of hard- which is now a state standard for etation Mapping Plan in the Mount wood trees in each stand, photo in- vegetation mapping in California. Tamalpais Watershed of southern terpreters evaluated how much of The summer of 2015 marked the Marin County, in part to address the existing trees (mainly tanoak or start of the 7th triennial vegetation wildfire risk and the severity of Sud- coast live oak) in the canopy died survey of Suisun Marsh and was con- den Oak Death (SOD) caused by over time. Updates to the original ducted by the Vegetation Classifica- the plant pathogen Phytophthora 2004-baseline map highlighted ad- tion and Mapping Program within ramorum. The original baseline veg- ditional SOD affected areas, quanti- the California Department of Fish etation map, created in 1993 by East- fied hardwood cover loss created and Wildlife. West Forestry Associates noted from previous die-offs, and captured Suisun Marsh is located in Solano tanoak as the most abundant tree, floristic change caused by the re- County and is part of the San Fran- two years prior to the arrival of SOD. sultant hardwood death. The most cisco Bay/Sacramento–San Joaquin In 2004, Aerial Information Systems recent effort, conducted in 2014, River Delta estuary ecosystem. It is Inc. (AIS) created a vegetation map goes a step further by evaluating all one of the largest contiguous brack- using modern imagery and the MCV polygons containing hardwoods ish marshes remaining in the United classification system through a pro- (roughly 13,000 acres) and individu- States, covering almost 70,000 acres cess known as heads up digitizing. ally quantifying the total area of of tidal and managed seasonal wet- Their map was updated in the spring canopy openings created by SOD. lands. The amount of information of 2009 and again in 2014, so that This effort revealed that the wa- that can be gleaned from such a over a ten-year period, photo inter- tershed has lost nearly all its exist- long, on-going, fine scale mapping preters from AIS produced three ing stands of tanoak forest, in addi- project can make your head spin. vegetation maps that spatially rep- tion to suffering a serious decline in Examples include: detecting and

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 23

V45_2_book.pmd 23 6/9/17, 6:53 AM monitoring changes in preferred standard conventions, a vegetation Regional Conservation Authority. habitat for the endangered Salt map can be updated and compared Redlands, CA. Marsh Harvest Mouse, tracking the to earlier versions, depicting and Aerial Information Systems, Inc. (AIS). spread of non-native species of quantifying changes to the struc- 2015c. Summary report for the 2014 concern including common reed ture, composition, quantity, and photo interpretation and floristic (Phragmites australis) and perennial quality of the patterns of vegeta- reclassification of Mt. Tamalpais pepperweed (Lepidium latifolium), tion. Because vegetation mapping watershed forest and woodlands project. Prepared for the Marin Mu- correlating management activities efficiently and accurately represents nicipal Water District. Redlands, CA. with vegetation changes, providing plant distribution across broad ar- California Department of Forestry and baseline data for current and future eas it is a very important tool for Fire Protection. 2014. Fire and Re- restoration and climate change stud- understanding present, past, or fu- source Assessment Program (FRAP). ies, and much more. ture conditions in larger areas than Fire perimeters geodatabase. Common reed is a species of con- can be efficiently tracked by tradi- frap.fire.ca.gov/data/frapgisdata-sw- cern because it is wide-spread, it tional individual plant species moni- fireperimeters_ download.php displaces native species, and it has toring methods. Jones and Stokes Associates, Inc. 1993. rapidly increased in Suisun Marsh The accessibility of high quality Methods used to survey the vegeta- since 1999 (Figure 5). When com- aerial imagery and the availability of tion of Orange County parks and paring vegetation maps over time, vegetation maps online makes com- open space areas and the Irvine com- common reed has increased by 216% parison over time more feasible now pany property. Unpublished report over the entire marsh, starting with than ever before. A timeline of aerial prepared for County of Orange, En- only 863 acres in 1999 and covering imagery is currently available online vironmental Management Agency. 3,142 acres by 2015 (VegCAMP in through Google Earth, allowing us- Santa Ana, CA. prep). Vegetation maps are an effec- ers to look back in time at various Kelly, M., B. Allen Diaz, and N. Kobzina. 2005. Digitization of a his- tive tool to show where species like landscapes. Repeat vegetation map- toric dataset: the Wieslander Califor- common reed are increasing rapidly ping can show stability, increase, or nia vegetation type mapping project. or displacing preferred habitat for loss of certain species of plants or Madroño 52(3):191–201. the Salt Marsh Harvest Mouse, in plant groups. Taken at face value, Kuchler, A.W. 1977. The map of natu- order to minimize losses and priori- these observations may be only in- ral vegetation of California. tize control. teresting to botanists or biogeogra- Thorne J.H., B.J. Morgan, and J.A. Suisun Marsh is an important re- phers, however, when taken in con- Kennedy. 2008. Vegetation change source that is owned by several agen- text of cause and effect, these obser- over sixty years in the central Sierra cies and many private entities but vations can be strong support for Nevada, California, USA. Madroño. the management is highly regulated informed conservation or manage- 55(3):223–237. and treated as one unit. A compre- ment decisions about individual Vegetation Classification and Mapping hensive Suisun Marsh Habitat Man- species, natural communities, or Program (VegCAMP). In prep. 2015 agement, Preservation, and Resto- ecoregions. vegetation map update for Suisun ration Plan was completed in 2014. Marsh, Solano County, California. A report to the California Department According to this plan, the next 30 REFERENCES years will bring over 5,000 acres of of Water Resources. California De- tidal marsh restoration and over Aerial Information Systems, Inc. (AIS). partment of Fish and Wildlife, Sac- ramento, CA. 40,000 acres of managed wetland 2005. 2001 Western Riverside Wieslander, A.E. 1935. A vegetation enhancement across Suisun Marsh. County vegetation map. Prepared for the Western Riverside County type map of California. Madroño. Fine scale vegetation maps are cru- 3:140–144. cial for tracking the progression of Regional Conservation Authority. Redlands, CA Wright, D., C.V. Nguyen, and S. Ander- the vegetation and channel mor- son. 2016. Upward shifts in recruit- phology for these restored and en- Aerial Information Systems, Inc. (AIS). 2015a. Orange County vegetation ment of high-elevation tree species hanced sites. mapping update phase II. Final in the northern Sierra Nevada, Cali- vegetation mapping report. Prepared fornia. California Fish and Game Jour- SUMMARY for the Nature Reserve of Orange nal. 102(1):17– 31. County. Redlands, CA. Every member of CNPS knows Aerial Information Systems, Inc. (AIS). Todd Keeler-Wolf, Todd.Keeler-Wolf@ and appreciates California’s great 2015b. Western Riverside County wildlife.ca.gov; Rachelle Boul, Rachelle_ floristic diversity and the sensitivity vegetation mapping update. Final [email protected]; John Menke, jmenke of our flora to both natural and hu- vegetation mapping report. Prepared @aisgis.com; Jennifer Buck-Diaz, jbuck@ man agents of change. By following for the Western Riverside County cnps.org

24 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 24 6/9/17, 6:53 AM Great Valley phacelia (Phacelia ciliata), common monolopia (Monolopia lanceolata), and California poppy (Eschscholzia californica) in the Tehachapi foothills on the Antelope Valley side of Tejon. Photograph by Nick Jensen. TEJON RANCH—LAND OF CONTRAST, BOTANICAL RICHNESS, AND ONGOING DISCOVERY by C. Ellery Mayence, Nick Jensen, Neal Kramer, Laura Pavliscak, and Michael D. White

TEJON RANCH— and Los Angeles Counties, just over “when California was wild, it was AN INTRODUCTION an hour’s drive north of the Los An- the floweriest part of the continent.” geles Metropolitan area. Remarkably, For that matter, experiencing bo- ome have heard of it, many this landscape represents 0.25% of tanical displays on the Ranch even have driven past it, but few California’s land area yet supports in a so-called ‘drought’ year can still have experienced the natu- 13% of the state’s native flora. Expe- incite a lengthy list of superlatives. ral wonder of Tejon Ranch. riencing Tejon in a year flush with There is no better way to convey STejon Ranch, California’s largest precipitation when wildflowers are the ecological significance of Tejon contiguous private property at in full bloom embodies a landscape than through a discussion of its bo- 270,000 acres, is located in Kern indicative of John Muir’s words tanical resources—resources that

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 25

V45_2_book.pmd 25 6/9/17, 6:53 AM have yet to be completely censused of special concern (e.g., California as evidenced by a plethora of recent jewelflower, Caulanthus californicus, significant botanical discoveries that Rank 1B.1; San Joaquin adobe sun- by all indications will continue as burst, Pseudobahia peirsonii, Rank more surveys are conducted. As ex- 1B.1; Fort Tejon woolly sunflower, pressed by Principe and White Eriophyllum lanatum var. hallii, Rank (2015), Tejon Ranch is one of the 1B.1), as well as at least two species most prominent hidden treasures of endemic to the Ranch, Tehachapi the Tehachapi region. Our objective buckwheat (Eriogonum callistum, here is to comment on how and why Rank 1B.1) and Comanche Point Tejon Ranch supports such botani- layia (Layia leucopappa, Rank 1B.1). cal diversity, convey the overall im- Describing Tejon’s rare plant com- portance of conducting floristic sur- munities in detail is a manuscript in veys on the Ranch, and describe how itself and beyond the scope of this they influence land management. piece. Equally as fascinating botani- We provide examples of botanical Tejon Ranch is one of the most prominent cally and an indication of how geog- discoveries within interesting eco- hidden treasures of the Tehachapi Region— raphy and physiography influence and a significant piece of California’s biological contexts and convey the role diversity conservation puzzle. Map not species richness are the number of of the Tejon Ranch Conservancy in drawn to scale. taxa representing specific genera. ensuring the Ranch’s natural re- Source: C. Ellery Mayence, Tejon Ranch Buckwheat (Eriogonum spp.), for ex- sources are appropriately managed. Conservancy, 2016 ample, is ubiquitous across the Ranch and represented by an astounding WHY IS TEJON SO ley, Mojave Desert, and the south- 27 (and counting) taxa. Other gen- BOTANICALLY RICH? ern Sierra Nevada, Coast, Tehachapi, era represented by ten or more na- and Transverse mountain ranges. tive taxa include: Gilia (18), Lupinus There are many interconnected This equates to the Ranch represent- (14), Clarkia (14), Cryptantha (12), reasons as to why Tejon Ranch sup- ing what essentially is one large tran- Phacelia (12), Quercus (12), Juncus ports a large fraction of California’s sition zone between well-defined (10), and Elymus (10). More than 40 botanical diversity. Above all, the biomes with distinct floristic asso- genera are represented by five to nine Ranch sits at the convergence of sev- ciations. Tejon supports at least 45 native taxa. eral prominent physiographic re- taxa classified by the California Na- The Ranch is a rugged landscape gions including the San Joaquin Val- tive Plant Society (CNPS) as species with significant topographical relief,

Grassland habitat, a focus of the Conservancy’s land management efforts, comprises more than 100,000 acres of the Ranch and exists in different forms. LEFT: San Joaquin Valley grassland dominated by non-native annual Bromus and Avena species, and RIGHT: Antelope Valley grassland with a larger fraction of native bunchgrasses including Stipa speciosa. Photographs by C. Ellery Mayence.

26 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 26 6/9/17, 6:53 AM a diverse geology, and varied soil that is, the greater the land area the types, some of which are relatively greater the likelihood that more spe- uncommon elsewhere in the region. cies can be supported. Therefore, Elevation on Tejon ranges from a size does matter with respect to eco- few hundred feet above sea level to logical processes and the provision approximately 6,800 feet, rising over of habitat, specifically as it pertains a relatively short distance (<10 to the maintenance of local adapta- miles). Situated at the nexus of mari- tion, regional genetic diversity, and time and desert climates and influ- plant population viability. Globally, enced by topography, slope, and as environments become increas- aspect are a multitude of micro- ingly homogenized through habitat climates that, in part, drive plant fragmentation, extinction, and the diversity and community composi- proliferation of well-adapted, non- tion. A product of this diverse abi- native species, landscapes such as otic environment is the development Tejon take on an increasingly im- of unique and diverse soil forma- portant role in conserving bio- tions interspersed across the Ranch. diversity. The scale of Tejon Ranch Indeed, soil characteristics appear along with minimal outside influ- to be one of the single most impor- ence are fundamental to understand- tant drivers of plant diversity on ing how and why Tejon supports on Tejon. Principe and White (2015) the order of 1000 plant species. nicely describe the variety of special status species that inhabit the alka- BOTANICAL DISCOVERY, line sandy loams and deep clays of RARE PLANT REFUGIA, the northern and southern Tejon AND SIGNIFICANT RANGE Hills on the San Joaquin Valley side of the Ranch. Perhaps equally as EXTENSIONS Collections destined for the herbarium— botanically important, though sig- The more time spent afield, the a critical element in documenting and nificantly less inventoried, are meta- greater the likelihood of new obser- understanding Tejon’s flora. Photograph by sedimentary carbonate outcrops on vations and discoveries. This is the C. Ellery Mayence. the Antelope Valley side of Tejon essence of botanizing on Tejon. Be- laid down in shallow inland seas and ing private property with restricted than five-fold to the 7,000 collec- uplifted with the Tehachapi Moun- public access means that compared tions in existence today. tains. Well known as the only habi- to more accessible public and pri- If one factors in yearly and even tat of Eriogonum callistum, it is al- vate land, Tejon’s botanical re- decadal variation in weather and cli- most a given that additional note- sources as a whole have yet to be mate, soil type, specific cues affect- worthy discoveries await in this area. as thoroughly cataloged. Botanical ing seed germination and plant re- The volcanic soils straddling the San surveys and discoveries increased cruitment, and phenological differ- Andres rift valley are another edaphi- significantly eight years ago with ences across species, it is easy to not cally unique region of Tejon that has the creation of the Tejon Ranch observe certain species in a given been scantly explored by botanists. Conservancy who commissioned year as well as to rarely encounter As with discoveries elsewhere, ex- the work of consulting botanist species that, based on herbarium tensive surveys will need to be con- David Magney, followed by survey records, should be present. This ducted in years with optimum pre- work by consulting botanist Neal could happen for a number of rea- cipitation and temperature regimes— Kramer and Conservancy employ- sons: (1) conditions were not con- two factors not so easily manipu- ees. Just as significant is botanist ducive for germination that year, (2) lated. Although unexpected for an Nick Jensen’s ongoing, multi-year germinants and/or seedlings were area in close proximity to one of the effort to document and write the predated or died early in develop- most populous parts of the state, the first ever flora of Tejon. Prior to the ment and did not mature, (3) sur- black box of botanical discovery on creation of the Conservancy and veys were not conducted in suitable Tejon awaits further exploration. stemming from surveys associated habitat and/or at the appropriate It is generally accepted that a with proposed development proj- time, or (4) populations were too positive relationship exists between ects, about 1,300 botanical collec- small and inconspicuous to be land area and species diversity tions were known from the Ranch. readily observed in such a large, to- (MacArthur and Wilson, 1967)— This number has increased more pographically complex setting. The

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 27

V45_2_book.pmd 27 6/9/17, 6:53 AM Tejon supports at least 45 special status plant species. Shown are: a) California jewel flower (Caulanthus californicus), b) Peirson’s lupine (Lupinus peirsonii), c) Lost Hills crownscale (Atriplex coronata var. vallicola), and d) San Gabriel manzanita (Arctostaphylos glandulosa subsp. gabrielensis). Photographs by Neal Kramer.

Tejon Hills are a hotspot of rare observed. Whether isolated sites in 1B.2), and Mount Gleason paint- plant occurrences and certainly a the Tejon Hills serve as refugia for brush (Castilleja gleasoni, Rank major destination for rare plant en- rare species previously widespread 1B.2) are CNPS special status spe- thusiasts. Though special status spe- is not known, though it is interest- cies previously considered endemic cies were known to inhabit the Tejon ing that other locations with similar to the San Gabriel Mountains or Hills as early as 2011 if not before, it characteristics (e.g., elevation, soil further afield yet now known to was not until the winter of 2015-16 type) do not support the same suite occur on Tejon Ranch. Such dis- when precipitation amount and tim- of species, or at least have not been junct distributions bring to light ing were more typical for the region observed to do so. several interesting biogeographical after several years of drought that Range extensions add another questions, specifically whether such numerous rare species were ob- level of complexity and intrigue to species were more widespread and served in significantly large num- rare plant distributions, especially have retreated to their current dis- bers. Aside from surveying inten- when over long distances and/or tributions as a result of climate sity, phenology varies considerably across significant geographical bar- change over the past several thou- across species so unless target areas riers. For example, Peirson’s lupine sand years. Alternatively, it is pos- are routinely surveyed, it is possible (Lupinus peirsonii, Rank 1B.3), San sible for seed to disperse over long for fast-growing, annual species to Gabriel manzanita (Arctostaphylos distances, with the observed indi- complete their life cycle and not be glandulosa subsp. gabrielensis, Rank viduals representing founder populations. Botanical discovery that results in significant range extensions provides clues into broader biogeographical processes that may not be apparent under current environmental conditions.

PLANT CONSERVATION AND LAND MANAGEMENT Working landscapes, notably cattle ranches like Tejon, elicit a different conservation and land management framework compared to lands managed solely for low-im- With so many species comes the full spectrum of colors and variants. Shown are: (left) pact recreation and conservation. sapphire woollystar (Eriastrum sapphirinum) and (right) San Joaquin adobe sunburst Simply identifying a new popula- (Pseudobahia peirsonii). Photographs by Neal Kramer. tion of a special status species does

28 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 28 6/9/17, 6:53 AM not necessarily equate to a change that benefit their persistence, albeit vancy does hinges on scientific re- in land management, but rather, at on a small spatial scale. For example, search, it collaborates with educa- least in the short term, greater devo- when a population of California tional institutions on all levels and tion to stewardship and monitoring jewel flower (C. californicus) was provides logistical support for nu- to assess population structure and discovered in the Tejon Hills, vol- merous Master’s and Doctoral re- change over time. This is particu- unteers were organized to physically search projects. larly true on Tejon where cattle have remove Saharan mustard (Brassica been part of the landscape since the tournefortii) from the vicinity. Simi- LOOKING AHEAD mid-19th Century. Many of the plant larly, when a population of western species known to occur on the Ranch dog violet (V. adunca subsp. adunca) The future is bright for the con- today, special status species in- was discovered, the first for this spe- servation of botanical resources on cluded, have persisted despite con- cies in Kern County, it was deter- Tejon Ranch and it is likely that sistent grazing and trampling by live- mined that to meet its habitat re- forthcoming botanical surveys will stock—though historical distribu- quirements it would be necessary to yield new species, as such discovery tions for most are poorly understood. limit encroachment by nearby wil- shows no sign of slowing. Of the Though it goes without saying lows. As such, conservation and land property’s 270,000 acres, approxi- that Tejon likely supported greater management approaches will differ mately 88% or 240,000 acres are native botanical diversity prior to based on species requirements and protected under an extensive net- European settlement, one could ar- site-specific attributes and should work of conservation easements. As gue that being a working landscape be adaptable to changing conditions. has been the case since it was cre- for 100+ years may have actually ated, the Conservancy will continue benefited conservation of species not THE ROLE OF THE to use in-situ research and best avail- initially lost as land utilized by na- CONSERVANCY able science to achieve conservation tive peoples was consumed by in- goals and enhance land manage- tensive agrarian uses. This is not to The Tejon Ranch Conservancy, a ment. As opportunities arise, the say that grazing has resulted in an 501c3 not-for-profit accredited land condition of degraded habitats will increase in plant diversity or is ben- trust, was created in 2008 and works be improved with the restoration of eficial in all cases, but rather that to protect, enhance, and restore the key ecosystem processes and ser- persistent grazing has likely helped native biodiversity and ecosystem vices serving as success criteria. offset complete transformation by values of Tejon Ranch and the non-native Mediterranean grasses Tehachapi Range for the benefit of of, for example, the San Joaquin California’s future generations. In REFERENCES Valley side of the Ranch. These in- doing so the Conservancy works Bartolome, J.B., B. Allen-Diaz, S. Barry, vasive grasses are superior competi- closely with the land owner, the I. Ford, M. Hammond, P. Hopkinson, tors compared to many native Cali- Tejon Ranch Company, to prioritize F. Ratcliff, S. Spiegal, and M.D. fornia forbs and grasses (Bartolome and implement conservation and White. 2014. Grazing for biodiver- et al., 2014), and can radically alter land management activities. To sity in Californian Mediterranean landscape-scale ecological pro- achieve its internal objectives, the grasslands. Rangelands 36(5):36–43. cesses. When implemented prop- Conservancy has established three MacArthur, R.H., and E.O. Wilson. erly, grazing can decrease the com- program areas: Science, Stewardship, 1967. The Theory of Island Biogeog- petitive edge held by non-native spe- and Public Access and Education. raphy. Princeton University Press. cies and minimize thatch accumu- These programs operate interactively Principe, Z., and M.D. White. 2015. lation, both of which benefit native with university and Conservancy- Hidden treasures of the Tehachapi plant persistence. However, the ben- led research projects, citizen science, Region. Fremontia 43(2):3–9. efit of grazing is very much site de- stewardship projects, conservation pendent—and debatable in some easement monitoring, environmen- C. Ellery Mayence, Laura Pavliscak, and cases. For land managers, the ques- tal education and outreach, and pri- Mike D. White are affiliated with the tion becomes when, where, and at vate and public outings represent- Tejon Ranch Conservancy, PO Box 216, what intensity is grazing benefi- ing major activities. For native plant Frazier Park, CA 93225, emayence@ cial—or not? enthusiasts, the Conservancy hosts tejonconservancy.org; Nick Jensen, Rancho As new populations of special sta- numerous CNPS chapters through- Santa Ana Botanic Garden, 1500 North tus species are identified, effort is out the year and collaborates with College Avenue, Claremont, CA 91711, made, especially for extremely small the Jepson Herbarium on at least [email protected]; Neal Kramer, Kramer populations or populations of very one multi-day workshop per year. Botanical, PO Box 1582, El Granada, CA rare species, to implement activities Because much of what the Conser- 94018, [email protected]

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 29

V45_2_book.pmd 29 6/9/17, 6:53 AM An exquisite stand of bigcone Douglas-fir looking toward Mount Markham from the Mount Wilson Toll Road, Angeles National Forest. Photograph by Michael Kauffmann. BIGCONE DOUGLAS-FIR MAPPING AND INVENTORYING IN THE ANGELES NATIONAL FOREST by Michael Kauffmann, Julie Evens, and Jaime Ratchford

he genus Pseudotsuga has a Douglas-fir appears in the fossil carpa (Axelrod 1961). Undoubtedly, present-day range that is record for the first time in the late the range of at least P. menziesii quite discontinuous. The spe- Pliocene, about 3 MA (Axelrod underwent repeated changes—cor- cies within the genus are 1950). The character of the fossil responding to climate trends— Tconfined to western North America flora accompanying bigcone Dou- throughout the Pleistocene across and eastern Asia. Unlike the exten- glas-fir was similar to today’s flora both the southern Coast Ranges and sive, continuous range of the more in the Transverse Range. This sug- southern Sierra Nevada. During this common Douglas-fir (Pseudotsuga gests that bigcone Douglas-fir may time it is possible that the two menziesii) across the western United have always been restricted in range species repeatedly overlapped in States, bigcone Douglas-fir (Pseudo- (Hermann 1985). More recent fossil range—though, like today, never hy- tsuga macrocarpa) is a Southern Cali- evidence indicates that in the early bridized. fornia endemic (Figure 1). Pleistocene, a time that was cooler Asian Pseudotsuga species and P. and wetter, the forest flora that macrocarpa are the more ancient spe- PSEUDOTSUGA included P. menziesii extended fur- cies within the genus while the mod- ther south than it does today, possi- ern P. menziesii most likely evolved The fossil record for the genus bly into what is now Baja Califor- during the Pleistocene. Since that begins in the early Tertiary, about nia, and inhabiting the Transverse time, P. menziesii has gone on to 50 million years ago (MA). Bigcone Ranges concurrently with P. macro- establish itself as a major compo-

30 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 30 6/9/17, 6:53 AM nent of the forest vegetation in western North America, as well as one of the most important lumber trees in the world, while P. macrocarpa con- tinues to survive within a highly restricted range. While considered to have little economic value, ecosystem services include shade creation, forage and habitat for wildlife, and aesthetic beauty.

A NATURAL HISTORY Bigcone Douglas-fir spans the Transverse and Peninsular ranges— an extent of 135 miles from north to south and 210 miles east to west— across Santa Barbara, Ventura, Kern, Los Angeles, San Bernardino, Or- ange, Riverside, and San Diego counties. Northern range limits include areas around Mount Pinos in Kern FIGURE 1: Range of Pseudotsuga across the southwestern United States. From Kauffmann County and the headwaters of La 2013. Brea Creek in Santa Barbara County. Western limits include Zaca Peak in Village, was measured in 2005 to live oak (Quercus chrysolepis) forms the San Rafael Mountains. The 50m tall and 7m in circumference- the mid-canopy. Trees occur on southern and eastern limits are east making it one of the largest trees in gravelly soils that are usually shal- of Julian along Highway 78 in San the region. low, often with a sparse understory Diego County. Bigcone Douglas-fir inhabits a of Mediterranean-type chaparral Across this region the species zone generally between the lower species. ranges in elevation from 300-2,100 elevation chaparral and the higher Bigcone Douglas-fir is distinct m (1,000-7,000 ft) where it gener- elevation mixed-conifer forest. from other species in the genus be- ally occurs on cooler, north facing Within the chaparral slopes at lower elevations shifting to zone, the species is often south facing slopes in higher eleva- found in small patches. In tions. The most vigorous stands oc- mature forests, bigcone cur on the north slopes, which re- Douglas-fir forms the up- tain more moisture and lower aver- per canopy while canyon age temperatures. Habitats include chaparral, riparian canyons, mixed-conifer woodlands, and desert slopes, all of which offer rough terrain and steep aspects that are often very steep (over 50 ). The tall stature of the species, compared to many of the shrubs with which it associates, makes it quite distinct, especially large specimens. The two largest documented trees LEFT: Bigcone Douglas-fir seed cone (left) compared to coast Douglas-fir. • RIGHT: The second largest grow in the San Gabriel documented bigcone Douglas-fir was found during our surveys. It is 118' tall, with a 265" Mountains. The largest re- circumference, and a 114' crown spread for 412 total points (according to the American Forests Tree corded, near Mount Baldy Measuring Guidelines). Photo by Michael Kauffmann with Daniel Hastings.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 31

V45_2_book.pmd 31 6/9/17, 6:53 AM cause it has uniquely large cones has unusually thick bark—15-20 cm in the Sierra Pelona Mountains to and seeds (Table 1). With an in- in mature trees. Within one to three the northwest. These two popula- crease in cone and seed size, a func- years after fire events, buds not killed tions have quite distinct ecologies. tional shift in the mode of seed dis- will typically sprout new shoots Stands in the San Gabriel Moun- persal has occurred. Species that (Vander Wall et al. 2006). These ad- tains are in four general regions clas- have evolved characteristics such as aptations suggest bigcone Douglas- sified in a transect from the front well-defended cones and larger seeds fir has evolved to survive multiple range to the Mojave Desert (Figure typically have a closer relationship fire events that arrive via the sur- 3). The lowest elevation sites occur with animals as seed dispersers rounding chaparral with a short fire in the front-range, generally in north- (Vander Wall et al. 2006). Exclud- return interval (Lombardo et al. facing canyons or on steep slopes. ing Pseudotsuga macrocarpa, ances- 2009). A fire scar chronology on 85 The interior slope sites include river trally and currently all other species trees from 15 sites dating back to drainages and higher elevation peaks in the genus have winged seed for 1600 across Los Padres National like the San Gabriel River and the wind dispersal. Although P. macro- Forest suggests that in this region a Tujunga Canyon regions. The higher carpa has a winged seed, the mass of 400 year old tree would, on average, elevation sites are mixed conifer the seed renders this wing largely experience 13 fires in its lifetime stands where the species overlaps ineffective for wind dispersal. Ani- (Lombardo et al. 2009). with montane conifers. The north- mal-dispersed pines live in drier en- slope, or Mojave slope, stands com- vironments and animal-dispersed prise interesting associations of P. macrocarpa lives in drier environ- BIGCONE DOUGLAS-FIR IN mixed-evergreen plants like the rare ments too. Thus we see convergent THE ANGELES NATIONAL Parry’s manzanita (Arctostaphylos trends: animal seed dispersal allows FOREST parrayana subsp. parrayana) and occupancy of drier sites. other desert flora. Bigcone Douglas-fir is one of the Across the Angeles National For- In the Sierra Pelona Mountains, most fire resistant and adapted coni- est, bigcone Douglas-fir occurs in the species occupies a small per- fers in the world, with the ability to some of the highest densities for centage of acreage for the entire for- survive frequent fires because of the anywhere in the species’ range (Fig- est. Here, bigcone Douglas-fir mixes prolific resprouting of new needles ure 2). While the heart of this range with black oak (Quercus kelloggii) and branches from meristematic tis- is in the San Gabriel Mountains, a in open grasslands on the ridgeline sue on branches and boles. It also smaller, disjunct population occurs of Liebre Mountain down into the

FIGURE 2: Range of Pseudotsuga macrocarpa across the Angeles National Forest. From Kauffmann et al. 2017.

32 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 32 6/9/17, 6:53 AM Bigcone Douglas-fir stand on the south-facing slopes of the San Gabriel Mountain’s front range, overlooking the Inland Empire. Photo- graph by Michael Kauffmann

the north slope and canyons and included verifying distribution rapid assessments on bigcone Dou- into the upper elevations of the An- and status of bigcone Douglas- glas-fir stands. The data collected telope Valley near Neenach. Else- fir, ground-truthing polygons des- during field assessments were used where in the Sierra Pelonas, bigcone ignated by US Forest Service Re- to develop both an updated distri- Douglas-fir persists in small, iso- mote Sensing Lab’s California bution map via heads-up digitizing lated, relict stands on rocky sum- Vegetation (CALVEG) system as and a vegetation classification to mits or in sheltered north-facing “Bigcone Douglas-fir” Regional better define bigcone Douglas-fir river canyons. These low density Dominant, and conducting 370 stands for the area. stands persist because stand-replacing fires have not occurred in these small pockets. TABLE 1. CHARACTERISTICS OF CONES AND SEEDS OF DOUGLAS-FIR (PSUEDOTSUGA MENZIESII) AND MAPPING AND BIGCONE DOUGLAS-FIR (P. MACROCARPA)* INVENTORYING PROJECT Cone or seed trait P. menziesii P. macrocarpa The California Native Plant So- ciety, working in collaboration with Seed mass (mg) 24.8 ± 3.7 132.6 ± 23.8 the US Forest Service (USFS), initi- Seed length (mm) 7.3 ± 0.7 10.9 ± 1.0 ated field surveys in the summer/ Wing area (mm2) 52.0 ± 14.1 97.3 ± 32.0 fall of 2015 to assess the extent and status of the California endemic Descent velocity (m/s) 1.28 ± 0.22 2.47 ± 0.62 bigcone Douglas-fir in the Angeles *Adapted from Vander Wall et al. 2006 National Forest. Goals in the field

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 33

V45_2_book.pmd 33 6/12/17, 11:00 AM Bigcone Douglas-fir and Parry’s manzanita vegetation type on calcareous soil at the edge of the Mojave Desert, San Gabriel Mountains. Photograph by Michael Kauffmann.

GENERAL PATTERNS Mountains with just under 6% in have been impacted by stand-replac- the Sierra Pelona Range (Table 2). ing fires within the last ten years. Several patterns emerged in our No stand-replacing fires were docu- Across both ranges we found some survey data. Of the 35,300 acres of mented via heads-up digitizing or evidence (i.e. char on bark) of fire bigcone Douglas-fir on the Angeles ground truthing in the Sierra Pelo- in 77% of the surveys. National Forest, over 94% of those nas while 10% of the acreage in the Mortality between 10 and 90% stands occur in the San Gabriel San Gabriel Mountains appears to was strikingly high, with 19% of

FIGURE 3: Ecological cross-section of the San Gabriel Mountains featuring Bigcone Douglas-fir distribution. From Kauffmann et al. 2017.

34 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 34 6/9/17, 6:53 AM TABLE 2: AREA OF BIGCONE DOUGLAS-FIR POPULATIONS BY FOREST REGION.

TABLE 3. THE FLORISTIC CLASSIFICATION OF ALLIANCES AND ASSOCIATIONS ORGANIZED ALPHABETICALLY BY LIFEFORM. Newly defined associations are designated with an asterisk (*). Com- plete table available at http://cnps.org/cnps/vegetation/pseudotsuga- stands exhibiting mortality (gener- macrocarpa. ally between 10 and 20%). This mor- ASSOCIATION # SURVEYS tality is occurring at lower eleva- Pseudotsuga macrocarpa (Alliance) 7 tions on both the south slopes of the San Gabriel Mountains and the Pseudotsuga macrocarpa – Quercus agrifolia 3 north-slopes of the Sierra Pelonas. Pseudotsuga macrocarpa – Quercus chrysolepis 54 Most likely this mortality is due to Pseudotsuga macrocarpa – Quercus chrysolepis – (Acer xerification. macrophyllum)* 14 Pseudotsuga macrocarpa – Quercus chrysolepis – Abies concolor – VEGETATION PATTERNS Pinus spp. (P. lambertiana, jeffreyi, ponderosa)* 17 Pseudotsuga macrocarpa – Quercus chrysolepis – mixed conifer / A classification analysis for this Cercocarpus ledifolius* 4 project resulted in 21 alliances and 39 associations where bigcone Dou- Pseudotsuga macrocarpa – Quercus chrysolepis – Pinus coulteri* 25 glas-fir exists in the Angeles NF. Pseudotsuga macrocarpa – Quercus chrysolepis – Pinus An alliance is a hierarchical classi- monophylla / Fremontodendron californicum* 14 fication unit of vegetation defined Pseudotsuga macrocarpa – Quercus chrysolepis – Quercus by the diagnostic species of the up- wislizeni / Arctostaphylos glandulosa* 6 permost canopy layer and contains Pseudotsuga macrocarpa – Quercus chrysolepis / Bromus one or more associations. Associa- diandrus* 14 tions define the vegetation in finer detail and reflect the variation of Pseudotsuga macrocarpa – Quercus chrysolepis / Ceanothus the vegetation at a more local level. integerrimus* 35 Eleven new associations were clas- Pseudotsuga macrocarpa – Quercus chrysolepis / Cercocarpus sified in the Bigcone Douglas-fir montanus* 17 Forest Alliance (indicated by an as- Pseudotsuga macrocarpa – Quercus chrysolepis / Hesperoyucca terisk in Table 3) as a result of the whipplei* 14 fieldwork. Of the 35,300 acres mapped in Pseudotsuga macrocarpa – Quercus kelloggii* 5

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 35

V45_2_book.pmd 35 6/9/17, 6:53 AM the Angeles NF, 18,573 acres were alliance map identified 9,236 acres fore the 2009 Station Fire, which mapped as Bigcone Douglas-fir as Bigcone Douglas-fir Forest that resulted in some stand-replacement Forest Alliance. In contrast, the were not mapped previously in the of conifer vegetation in the land- CALVEG map had 34,055 acres at- CALVEG map. We suppose that the scape (e.g., canyon live oak, grass- tributed as bigcone Douglas-fir domi- resulting alliance map benefited from land, and chaparral vegetation). nant stands (USDA 2014). The dif- significant ground-truthing during ference between these two maps ap- the project, and from using more MANAGEMENT pears partially as an error of omis- detailed aerial imagery than was used RECOMMENDATIONS sion in the CALVEG map, since the in the existing CALVEG map of the region. Conversely, Because bigcone Douglas-fir there were 24,766 grow in two ecologically distinct acres mapped in the regions of the Angeles NF we pro- CALVEG map as vide recommendations for each re- bigcone Douglas-fir gion. See Kauffmann et al. (2017) dominant stands that for a more detailed account of rec- were not mapped as ommendations; below are some such in the new alli- highlights. ance map—this dif- ference in acreage is Sierra Pelona Mountains likely due in part to Thickets of conifers, predomi- the CALVEG map nantly bigcone Douglas-fir and being produced be- some Coulter pine (Pinus coulteri),

LEFT: Fuels management may have saved this stand of PSMA after the 2009 Sheep Fire in Lone Pine Canyon. Photograph by Michael Kauffmann. • BELOW: The Station Fire footprint extends up the slopes of Bear Canyon (near Mount Wilson) toward Tom Sloane Saddle where the fire stopped, leaving unburned forest on the left (NE facing) side of the picture. Photo by Michael Kauffmann.

36 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 36 6/9/17, 6:53 AM Bigcone Douglas-fir encroaching into a black oak woodland in the Sierra Pelona Mountains. View is to the north toward the Tehachapi Mountains. Photograph by Michael Kauffmann.

have been recruiting in open spaces quency increases with proximity to of black oak woodlands that were roads—many of which were easier formerly managed by the Serrano to construct through this western Indians (Yuhaviatam) as a food region of the mountains as compared source for acorns as well as open to the steeper, central and eastern hunting grounds (Keeley 2002). We portions. In addition, roads vector recommend using targeted pre- invasive species that are ignitable. scribed fire or conifer sapling re- While roads may increase fire moval by hand within the upper frequency, they may benefit big- elevation oak woodland habitat to cone Douglas-fir recruitment by ensure this habitat does not type- creating open space for seedling convert to conifer forest. establishment. Recruitment was high on naturally and anthropogenically San Gabriel Mountains disturbed slopes where there is The San Gabriel Mountains have newly-exposed mineral soil with proportionately less low-level mor- little to no competition from other tality than the Sierra Pelonas but plant species (Figure 4). We recom- higher incidence of stand-replacing mend the Forest Service disperse (last 10 years) fire events—mostly locally collected seeds in roadcuts due to the Station Fire. Fire fre- or on new landslides between 1,200-

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 37

V45_2_book.pmd 37 6/9/17, 6:53 AM 38 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 38 6/9/17, 6:54 AM FACING PAGE, TOP: Bigcone Douglas-fir in the San Gabriel Wilderness. Photos by Michael Kauffmann. • INSET: Daniel Hastings and Joslyn Curtis surveying for trees near Wrightwood, California. • FACING PAGE, BOTTOM: Low elevation mortality in bigcone Douglas-fir stands in the San Dimas Experimental Forest. • ABOVE: Bigcone Douglas-fir growing along the San Gabriel River on rocky slopes approaching 90 degrees.

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 39

V45_2_book.pmd 39 6/9/17, 6:54 AM FIGURE 4: Landscape-scale patterns in bigcone Douglas-fir seedling recruitment can be seen in this unique location near Crystal Lake. Graphic by Michael Kauffmann.

40 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 40 6/9/17, 6:54 AM 3,100 m (4,000-7,000 feet). This management approach will repopu- late and stabilize slopes and promote persistence of bigcone Dou- glas-fir. Other recommendations include environmental education and public outreach to promote fire safety and noxious weed awareness. Away from roads, recruitment appears to be most common in wilderness areas that have not been managed due to their remote nature. With lack of management, fires have remained unconfined, of generally View down Lone Pine Canyon toward Victorville, Caifornia. Photo by Michael Kauffmann. low-intensity, and frequent. These historical fire regimes are important mon part of the species’ ecology, Kauffmann, M.E. 2013. Conifers of the to maintain because they are the con- allowing them to continue to flour- Pacific Slope. Backcountry Press. ditions to which this species has ish, for now, across the Angeles Na- Kneeland, California. adapted over the past five million tional Forest. Kauffmann, M., J. Ratchford, J. Evens, years. Steep slopes in the interior K. Lindke, and J. Barnes. 2017. An- geles National Forest: Bigcone Dou- mountains have offered long-term ACKNOWLEDGMENTS refugia from high-intensity stand- glas-fir mapping and monitoring report. Unpublished report. Califor- replacing fire. Recommendations in CNPS fieldwork staff included nia Native Plant Society Vegetation remote areas of the forest include Daniel Hastings and Joslyn Curtis. Program, Sacramento, CA. adding wilderness designation and Angeles National Forest Fuels Plan- Keeley, J.E. 2002. Native American eliminating roads where possible. ner Diane Travis, Southern Province impacts on fire regimes of the Cali- Ecologist Nicole Molinari, Humboldt fornia coastal ranges. Journal of Bio- THE FUTURE State University professor Lucy geography, 29: 303–320. Kerhoulas, and Anton Jackson with Lombardo, K.J., T.W. Swetnam, C.H. Across the forests of the Trans- TEAMS Enterprise Unit. Their sup- Baisan, and M.I. Borchert. 2009. Us- verse and Peninsular Ranges, high port in project funding and review ing bigcone Douglas-fir fire scars and potentials for stress are imminent in of the article are greatly appreciated. tree rings to reconstruct interior certain portions of the species range chaparral fire history. Fire Ecology due to climate-water deficits and 5(3): 32-53. REFERENCES large-scale fires. Bigcone Douglas- USDA Forest Service (USFS). 2014. Vegetation mapping. Existing veg- fir is likely to find refugia along the Axelrod, D.I. 1950. Further studies of etation (eveg) layers. Pacific South- higher, north facing elevations in the Mount Eden flora, southern Cali- west Region Remote Sensing Lab. the Angeles NF and potentially fur- fornia. Contributions Paleontology Data available at: http://www.fs.usda. III: 75-117. Carnegie Institution of ther north through the Tehachapi gov/detail/r5/landmanagement/ Washington Publications. Mountains, Sierra Nevada, and Coast resourcemanagement/?cid=stelprdb Axelrod, D.I., and W.S. Ting. 1961. Range over the next 85+ years. 5347192 Early Pleistocene floras from the Even before the current human- Vander Wall, S.B, M.I. Borchert, and Chagoopa surface, Southern Sierra induced climate changes, fossil evi- J.R. Gworek. 2006. Secondary seed Nevada. University of California dence suggests that the range of the dispersal of bigcone Douglas-fir Publications 39(2):119-194. species has been in decline. The seeds. Arcta Ecologica 30, 100-106. Gause, G.W. 1966. Silvical character- species is highly valued across the istics of bigcone Douglas-fir mountains of southern California (Pseudotsuga macrocarpa) [Vasey] FOR THE FULL REPORT as an endemic species, for its aes- Mayr). Berkeley, CA, Pacific SW. VISIT: thetic beauty, wildlife habitat, and Forest and Range Exp. Station. 10 influence on regional biodiversity pp., illus. (US Forest Serv. Res. Pa- cnps.org/vegetation/pseudotsuga- with the habitat it creates. These are per PSW-39). macrocarpa relict habitats where high severity Hermann, R.K. 1985. The genus Pseu- fires have been rare but other dis- dotsuga: Ancestral history and past Michael Kauffmann, mkauffmann@cnps. turbances like landslides and lower distribution. Forest Research Labo- org; Julie Evens, [email protected]; Jaime intensity fires have remained a com- ratory. Oregon State University. Ratchford, [email protected]

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 41

V45_2_book.pmd 41 6/9/17, 6:54 AM BOOK REVIEWS

Gathering Moss: A Natural and Cul- stabilizes her position as an unseen planted urban bee garden can attract tural History of Mosses by Robin Wall and unbiased observer-guide and in- 40 to 50 species of bees. Kimmerer. Oregon State University troduces the importance of a change As a beekeeper, this book is for the Press. $18.95. ISBN: 978-0870714993 in attention, something she invites the beekeeper and bee enthusiast alike, reader to do as well. Throughout the with reference to the social honeybee Robin Wall Kimmerer’s Gathering book, she situates some of her impor- and world of solitary native bee spe- Moss: A Natural and Cultural History of tant research questions in bryology into cies. It’s a book for the avid gardener Mosses guides readers through a shift the story of her life. By revealing the and flower enthusiast. I was interested in perspective towards what is over- personal side of scientific inquiry, she to learn more about specific bee-plant looked, under-valued or—at the ex- makes the questions all the more vivid interactions—for example the female treme—the completely un- for the reader. This personalized ele- squash bee (in the family Apidae) vis- seen. The book is a natural ment of the book lends it to being a its squash flowers in the early morn- history guide to mosses, brim- great read for young adults with an ing before sunrise, followed by a later ming with contextualized ana- interest in biology, plants, and/or natu- honeybee visit. When the flower closes tomical definitions and clearly ral history. in late morning you can peek inside presented ecological principles The blurriness of subject-object and often times find one or more male that incite a sense of wonder lines is itself, ultimately, the shift squash bees fast asleep-where it will for this small-scaled world. Kimmerer asks her readers to make. stay until the following morning! Woven throughout the narra- While Kimmerer may impose such Early chapters are loaded with de- tive, however, Kimmerer gen- clearly defined roles in the process of tailed information ranging from the tly nudges readers towards a more sub- scientific inquiry, in the other realms sophisticated lifecycle and bee identi- stantial shift in perspective, from na- of her life as a mother, a neighbor, an fication characteristics to equally so- ture-as-simple object to nature-as-in- indigenous woman, and a human be- telligent teacher. ing on an ecologically-threatened Each chapter is arranged as a vi- planet, mosses and nature have pow- gnette-length personal narrative and erful agency and wisdom from which tackles various aspects of moss biol- we can learn essential principles. A ogy and ecology, showing the reader beautiful chapter entitled “The Forest there is always more than meets the Gives Thanks to the Mosses” tracks eye. In one such vignette, Kimmerer how, in myriad ways, mosses are intri- takes time to explain the importance cately connected to the thriving of the of the boundary layer—the meeting forest ecosystem and its many mem- point of air and land—to mosses exist- bers. If Nature is our teacher, the main ence and relationship to water, all lesson is one of reciprocity and inter- to levels of detail that will satisfy avid connectedness. naturalists. Gathering Moss is a look at over- Other chapters look into mosses’ looked worlds: mosses, indigenous roles in ecological succession and res- ways of knowing, and Nature, to toration, variations in moss reproduc- name but a few. It merits more than a tive strategies, and moss communi- glance. ties. The author also introduces many —Catherine Curley of her favorite moss genera such as Dicranum, Dendroalsia, Tetraphis, Bryum, and Sphagnum—many of which California Bees and Blooms: A Guide have figured into her experiments. for Gardeners and Naturalists by Gor- phisticated bee mimicry and anatomi- Yet Kimmerer is not content to keep don W. Frankie, Robbin W. Thorp, cal disparities. Here we’re offered an the subject-object lines clear and one Rollin E. Coville, and Barbara Ertter. in-depth look at bee identification by of the main ways she confounds those Heyday. $28.00. ISBN: 978-1597142946 wing characteristics or other ways such lines is by turning the lens onto her- as host plants visited, seasonality, for- self. In the very first chapter, we ac- California Bees and Blooms opens in aging behavior and geographic distri- company her on a dreamy passage into homage to the return of John Muir’s bution. Chapters are then divided up a mossy clearing within a boulder pile, ‘bee pastures’ and an introduction to into bee families grouped by descrip- which she pointedly recounts, she has pollinator decline. Acknowledging we tors such as “sweat bees” (those that never seen before despite having can no longer rely on honeybees for lap up our perspiration!), mason bees walked past this section of land hun- agricultural pollination services-we (the architects) and mining bees. dreds of times. As such, Kimmerer de- look to the wild pollinators. One well- Within the chapters, bee profiles and

42 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 42 6/9/17, 6:54 AM close-up color photography detail the George Poinar has organized 22 most common genera found in ur- the chapters based on habitat ban California. Each genus is further types, starting with the kelp described by general appearance, spe- and driftwood communities, cial features, similar looking insects, and for the strand and dune distribution and abundance, flight sea- community, describes 58 plant son and behavior, preferred flowers, species and their associated and nest habitat. We also learn about organisms. natural enemies of bees-that little white Each species, from micro- crab spider we see in our flowers is not scopic nematodes to trees, has so innocent! an informative photograph (or Cultivators of gardens will enjoy two or three). George took all the chapter about pollination syn- the photos, and because of that dromes-a set of flower characteristics I feel there is a consistency in that can be used to identify its most presentation that allows better likely pollinator. Another chapter is understanding. Fifty-eight spe- dedicated to the plants that consis- cies of plants form chapters tently attract native bees. Each plant with a text introduction that profile includes descriptive informa- describes the life styles of the tion, flowering season, what resource various creatures that are as- it offers bees, the most frequent bee sociated with that plant. In- visitors, bee ecology and behavior, and sect species with multiple life gardening tips. You can read up on forms will often have an image that lavender bush or western redbud of both larvae and adults. For you have growing in your yard to find each community, a table is pro- out more about its particular pollina- vided showing the detrivores, tors. Knowing the Latin names and herbivores, predators, omnivores, para- world: “At night many [sand fleas] fall families of plants goes a long way in sites, and symbionts among the spe- victim to…rove beetles that crush them navigating through this section. cies described. in their mandibles and suck up their After reading this book I am ex- In general, little is known about body liquids.” We learn that carrion cited to see the bee world with new the organisms that coexist with plants. beetles eat strawberries, ergot is a fun- eyes, and challenge myself to look be- But George tells us that “The presence gus that infects dunes grasses, and yond my social honeybees. What I of ants explains why most insect her- which insects pollinate dune orchids. might have passed off as a bumblebee, bivores on giant vetch are concealed The writing is poetic at times, very I may now see is a carpenter bee. And in plant tissue.” He is graphic in his descriptive and packed with details. now I’m on the lookout for one of description of the events of the natural His in-depth knowledge of this hidden those many metallic-colored bees. The book closes with ‘recipes’ for design- ing bee habitats that serve diverse pur- poses, and lastly, ways to take action to protect native bees. It is a well- organized and informative body of work. You, too, will undoubtedly see the bee world with new eyes after reading through this thoughtful and descriptive guidebook. —Denise Newman

A Naturalist’s Guide to the Hidden World of Pacific Northwest Dunes by George Poinar Jr. Oregon State University Press. $24.95. ISBN 978-0- 87071-854-0

Finally, a field guide to dune habitats along the Pacific Coast that is based on wildlife relationships rather than taxonomy. Rather than having sections on plants, birds, lizards, insects, etc.,

VOL. 45, NOS. 1 & 2, JUNE 2017 FREMONTIA 43

V45_2_book.pmd 43 6/9/17, 7:31 AM world is unsurpassed, and his enthusi- that for the public to understand and entific names in the index as a useful asm for it is infectious. Each animal care about wildlife, they have to have cross-reference. species account includes common a vehicle to understand it. This book It is small enough to carry along on name and scientific name, order and is the first of its kind and I hope other a hike of any of the 21 trails high- family; plants are named by common authors start publishing guides in this lighted, but of sturdy enough paper to name, scientific name and family. The format. handle such use for many years. You characteristics, habitat and range are —Steve Hartman could even drop it in a stream and described for each species, along with probably not damage it too much, but a comment. Both native and non-na- I don’t recommend that. tive species are described. Tahoe’s Spectacular Wildflower Trails If you read and use this book, you The book is fully indexed and in- by Julie S. Carville. Mountain Gypsy will indeed learn a lot about plants, cludes a glossary of terms. You’ll need Press. $29.95 ISBN: 978-0-692-69818- and if you hike the trails with this the index if you want to find anything 1 book in hand, you will be rewarded because the plants are listed in no par- and enriched in the process. ticular order and by common name. This wildflower guide is a delight- —David Magney If I have anything to complain ful book with many very good photo- about, it is the title. There was no graphs and lots of interesting informa- reason to limit it to the “Northwest” tion about the wildflowers of the Lake dunes. Of the 432 species listings, Tahoe region. This well-written book 83% occur anywhere along the Pacific is 277 pages long and chock full of coast between Washington and Cali- photographs, 877 to be exact. fornia. This book will be very useful to It is arranged in four basic sections: anyone who visits coastal dunes in background types of information, trees, California. wildflowers (arranged by flower color), George Poinar Jr. is a truly modern and the plants by selected trails in the naturalist. He understood wildlife re- broad Tahoe region. This guide iden- lationships when he was a professor tifies the plants covered by common at UC Berkeley and I was a student names, with reference to a plant’s sci- illustrating scientific books and pa- entific name infrequently (but at least pers for him back in the 1970s. His once). The work with amber (The Quest for Life in photographs Amber, Addison Wesley, 1994 and The of the plants Amber Forest: A Reconstructions of a by flower Vanished World, Princeton, 1999, both color uses by George Poinar Jr. and Roberta only com- Poinar) required that he studied many mon names; forms of life—trying to identify pre- however, historic life trapped in rocks. But there is a George is even more, he is a natural- complete list ist-educator, with a strong sense of of both com- conservation and the understanding mon and sci-

Wildly diverse plants for both novice and nerd

72 Channing Way, Berkeley 94710 Open Thursday, Friday, and Saturday www.oaktownnativenursery.info

44 FREMONTIA VOL. 45, NOS. 1 & 2, JUNE 2017

V45_2_book.pmd 44 6/9/17, 6:54 AM WHAT IS A LICHEN? Join Today! NEWS FLASH: Two’s Company, Three’s a Lichen? It is a composite organism made CNPS member gifts allow us to promoteecent andscientific protect findingsCalifornia’s from native Spribille plants andet al. their (2016) habitats. change Gifts our up of fungus andare a photosynthesiz- tax-deductible minus the $12 of the total gift which goes toward publication of Fremontia. ing partner (photo-partner). Last Runderstanding of the lichen relationship. For over 140 years we summer❏ $1,500 a team Mariposa of lichenologists Lily ❏ an- $600 Benefactorthought that lichens❏ $300 were Patron made of a❏ fungus $100 Plant and Lover an alga or nounced❏ $75 their Family discovery that many❏ $75 Internationalcyanobacteria. or Library Spribille’s❏ $45 team Individual discovered❏ that $25 Limitedmany lichen Income species contain two types of fungi. The first type (the one we already knew lichens contain two types of fungi CORPORATE/ORGANIZATIONAL (see NEWS FLASH). The photo- about) is typically in the group ascomycetes—the sac fungi. The ❏ $2,500 10+ Employees ❏ $1,000 7-10 Employees ❏ $500 4-6 Employees ❏ $150 1-3 Employees partner can be either a green alga or previously unseen fungal partner is a newly discovered lineage of a cyanobacteria. In a few lichens, yeast in the group basidiomycetes—the club fungi, which include both typesNAME of photo-partners are gilled mushrooms. These yeasts appear to be responsible for producing some of the unique chemical compounds found in the lichen’s present.ADDRESS This oddity is interesting because these lichens represent a fungal tissue. There is still much more to learn about these yeasts and symbioticCITY relationship between three their role in the lichenSTATE relationship. ZIP kingdoms of life: fungi from King- dom ❏Fungi, Enclosed green is a checkalgae made from payable King- to CNPS Membership gift: dom ❏Protista, Charge my and gift tocyanobacteria ❏ Mastercard ❏ Visaon their ability to Addedexist ondonation bare of:rock Unlike plants, lichens have no from Kingdom Eubacteria (formerly in the baking desert or survive expo- barriers to protect themselves from Card Number TOTAL ENCLOSED: Kingdom Monera). Additionally, sure to the extremes of outer space. extreme conditions. They have no Exp. date like many other organisms, lichens However, they are actually quite sen- waxy cuticle or stomata to prevent Signature ❏ Enclosed is a matching gift form provided by have numerous species of bacteria sitive to environmental conditions. water from leaving or pollutants Phone my employer that live on and in them. Lichens are passive organisms and from entering the lichen. They don’t Email ❏ I would like information on planned giving The lichen relationship is an ex- they have few mechanisms to miti- have roots either. There is no active amplePlease of make a mutualistic your check payablesymbiotic to “CNPS” re- andgate send changes to: California in their Native surroundings. Plant Society, 2707uptake K of water or nutrients in li- lationship—theStreet, Suite 1, partners Sacramento, benefit CA from 95816-5130. Tel: (916) 447-2677; Fax: (916) 447-2727; livingcnps.org/membership together. The ; fungalWeb site: partners cnps.org.; Email: [email protected] provide general protection for the CONTRIBUTORSphoto-partner. This (continued includes from a back struc- cover) tural ‘home’ within which the photo- SUBMISSION C.partner Ellery lives. Mayence Chemical is the compoundsSenior Ecologist and Science Program Coordinator of INSTRUCTIONS the Tejon Ranch Conservancy. He holds a BA in Geography from UNC-Chapel Hill,found an inMEM fungal in Resource tissue are Ecology unpalat- from Duke University, and a PhD in Conserva- CNPS members and others are tionable Biologyor even (Restoration toxic to some Ecology critters, focus) from the University of New Orleans. invited to submit articles for pub- Rikkethereby Reese protecting Næsborg the has photo-part- a PhD in lichenology from Uppsala University in Swe- lication in Fremontia. If inter- den,ner fromwhere herbivory. she studied Other genetic chemical relationships among tiny, inconspicuous lichens. ested, please first send a short Shecompounds is currently in researchingfungal tissue lichens func- and bryophytes growing on large coast red- summary or outline of what woods.tion as ultraviolet filters which pro- you’d like to cover in your ar- Lauratect the Pavliscak alga or hascyanobacteria been employed from as Tejon Ranch Conservancy’s Stewardship ticle to the Fremontia editor at Managerthe sun’s sinceharmful 2014. rays. She In holds return, a BAthe in Environmental Studies from UC Santa [email protected]. Instruc- Cruzphoto-partner and a MS feedsin Natural the fungi Resources by pro- from the University of Arizona. tions for contributors can be Jaimeducing Ratchford sugars through is Associate photosynthe- Vegetation Ecologist with CNPS, where she assists found on the CNPS website, withsis. In surveying, addition classifying,to producing and sugars, mapping of vegetation in California. www.cnps.org, under Publica- Michaellichens Whitethat contain is the Conservation cyanobacteria Science Director of the Tejon Ranch Conser- tions/Fremontia. vancy(collectively and an ecologist called cyanolichens)with over 25 years of experience in California. Mike holds aalso BA transformin Ecology, atmospheric Behavior, and nitrogen Evolution from UC San Diego and a PhD from the Saninto Diego a form State that University the fungi and can UC use. Davis Ecology Joint Doctoral Program. Fremontia Editorial Advisory Board Living together as a lichen, the part- Jim Andre, Phyllis M. Faber, ners are able to colonize a much Holly Forbes, Naomi Fraga, Dan widerACKNOWLEDGMENTS range of habitats than if they wereThe on time their and own. effort by all authors and co-authors reviewing and re- Gluesenkamp, Brett Hall, David Lichensreviewing are their valuable manuscripts as biological was invaluable in making this issue into what Keil, Patrick McIntyre, Kara Moore, indicatorsit has become. because I wouldthey are also respon- like to thank all the reviewers including Lucy Pam Muick, Bart O’Brien, Liv Lichens can be categorized into three general growth forms, similar to how we group siveKerhoulas, to environmental Nicole Molinari, change. Steve Li- Sillett, Tom Carlberg, and a few anony- O’Keeffe, Roger Raiche, Teresa mous folks for graciously volunteering plantstheir time as herbs, as editorial shrubs, consultants.and trees. The lichen growthSholars, forms Dick are: Turner, 1) foliose= Mike leafy, Vasey with a chens give the general impression of distinct upper and lower surface; 2) fruticose= shrubby or hair-like; and 3) crustose= thin being tenacious little organisms based crust tightly attached to its substrate.

VOL. 44,45, NO.NOS. 3, 1 DECEMBER& 2, JUNE 20162017 FREMONTIA FREMONTIA 3

V45_2_cover.pmdV45_2_book.pmd 3 6/9/17, 11:086:52 AM AM California Native Plant Society Nonprofit Org. 2707 K Street, Suite 1 Sacramento, CA 95816-5130 U.S. Postage PAID

MLP

MIX Printed on sustainably harvested paper containing Paper from /J, responsible sources 10% post-consumer content, processed chlorine-free. FSC ® www.fsc.org FSC C021872

CONTRIBUTORS FROM THE EDITOR

Marie Antoine lives in Arcata and teaches a class on lichens he first three articles in this issue take us on a journey and bryophytes at Humboldt State University. She also works T into the small and beautiful world of lichens and bryo- alongside her husband, Professor Steve Sillett, in studying the phytes. Mosses, liverworts, and lichens are significant com- structure and function of the world’s tallest and largest trees. ponents of ecosystems across the state but are often over- Shelly Benson has been observing lichens from the Bay Area looked because of our eye-level connection to vascular to British Columbia for the past 18 years and is interested plants. A closer look (and a hand lens) reveals a window in using lichens as biological indicators of climate and air into the wonderful world of charismatic cryptograms. Shelly quality. Benson, Rikke Reese Næsborg, and Marie Antoine have Rachelle Boul is an environmental scientist with the Cali- contributed exquisite stories sharing their love and passion fornia Department of Fish and Wildlife. for these often-overlooked critters—taking us on a journey Jennifer Buck-Diaz is a vegetation ecologist with the Cali- to better understand them. fornia Native Plant Society where she surveys, classifes, and maps vegetation of California. Todd Keeler-Wolf, Rachelle Boul, Jennife Buck-Diaz, Julie Evens is the Vegetation Program Director of CNPS, where and John Menke tell the story of how vegetation mapping is she directs fine-scale vegetation mapping and classifcation changing using data and imagery from the past at various projects in support of A Manual of California Vegetation. intervals. Their analysis is depicting and quantifying changes Todd Keeler-Wolf is the senior vegetation ecologist with the of patterns of vegetation across the state to better under- California Department of Fish and Wildlife, where he pro- stand past, present, and future conditions. motes vegetation classifcation through vegetation sampling C. Ellery Mayence, Nick Jensen, Neal Kramer, Laura and mapping projects. Pavliscak, and Michael D. White revisit Tejon Ranch—a John Menke is a vegetation mapper with Aerial Information place many of us at CNPS hope becomes a National Park. Systems in Redlands, California. He has mapped vegetation The crossroads of the Sierra Nevada, Tehachapi Mountains, throughout the state since 1976. and Transverse Ranges is 0.25% of the state’s land area Nick Jensen is currently a PhD student at Rancho Santa Ana supporting 13% of the species in our native flora. They Botanic Garden and former CNPS Rare Plant Program Direc- share that diversity with us and the role of the Tejon Ranch tor. He has a BS Environmental Horticulture from UC Davis. Conservancy in protecting those resources. Michael Kauffmann lives in Humboldt County working as Lastly, Julie Evens, Jaime Ratchford and I share a CNPS an educator, author, and ecologist. In his spare time, he is vegetation program collaboration with the Angeles National establishing a 360-mile hiking trail through the Klamath Forest. For two years we mapped and inventoried the Cali- Mountains that celebrates the region’s biodiversity. He calls fornia endemic bigcone Douglas-fir and in the process broad- it the Bigfoot Trail. ened our understanding of the health and distribution of Neal Kramer, a self-employed consulting botanist with this stunning conifer in the San Gabriel and Sierra Pelona projects throughout California, has worked with the Tejon Ranch Conservancy documenting botanical resources on Tejon mountains of southern California. Ranch since 2011. He holds a BA in Botany from UC Berkeley It is truly a pleasure to be guiding Fremontia—enjoy! and a MS in Forest Ecology from the University of Idaho. —Michael Kauffmann

FREMONTIA ((continued on inside back cover)) VOL. 44, NO. 3, DECEMBER 2016

V45_2_cover.pmd 4 6/9/17, 6:51 AM