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VOL. 42, NO. 1 • JANUARY 2014 FREMONTIA JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

CALIFORNIA’SVOL. 42, NO. 1, JANUARY DESERTS, 2014 PARTPART 1:1: BIOLOGYBIOLOGY ANDAND FREMONTIA ECOLOGYECOLOGY CALIFORNIA NATIVE PLANT SOCIETY CNPS, 2707 K Street, Suite 1; Sacramento, CA 95816-5130 FREMONTIA Phone: (916) 447-CNPS (2677) Fax: (916) 447-2727 Web site: www.cnps.org Email: [email protected] VOL. 42, NO. 1, JANUARY 2014 Copyright © 2014 MEMBERSHIP Membership form located on inside back cover; California Native Plant Society dues include subscriptions to Fremontia and the CNPS Bulletin Bob Hass, Editor Mariposa Lily ...... $1,500 Family or Group ...... $75 Kara Moore, Managing Editor Benefactor ...... $600 International or Library ...... $75 James M. André, Patrick McIntyre, Patron ...... $300 Individual ...... $45 Advisors Plant Lover ...... $100 Student/Retired/Limited Income . $25 Beth Hansen-Winter, Designer CORPORATE/ORGANIZATIONAL Brad Jenkins, Cynthia Powell, and 10+ Employees ...... $2,500 4-6 Employees ...... $500 Mary Ann Showers, Proofreaders 7-10 Employees ...... $1,000 1-3 Employees ...... $150 CALIFORNIA NATIVE STAFF – SACRAMENTO CHAPTER COUNCIL PLANT SOCIETY Executive Director: Dan Gluesenkamp David Magney (Chair); Larry Levine Finance and Administration (Vice Chair); Marty Foltyn (Secretary) Dedicated to the Preservation of Manager: Cari Porter Alta Peak (Tulare): Joan Stewart Membership and Development the California Native Flora Bristlecone (Inyo-Mono): Coordinator: Stacey Flowerdew Steve McLaughlin The California Native Plant Society Conservation Program Director: (CNPS) is a statewide nonprofit organi- Greg Suba Channel Islands: David Magney zation dedicated to increasing the Rare Plant Botanist: Aaron Sims Dorothy King Young (Mendocino/ Sonoma Coast): Nancy Morin understanding and appreciation of Vegetation Program Director: East Bay: Bill Hunt California’s native plants, and to pre- Julie Evens Vegetation Ecologists: El Dorado: Sue Britting serving them and their natural habitats Kern County: Dorie Giragosian for future generations. Jennifer Buck-Diaz, Kendra Sikes Education Program Director: Los Angeles/Santa Monica Mtns: CNPS carries out its mission through Josie Crawford Betsey Landis science, conservation advocacy, educa- Horticulture Program Director: Marin County: Carolyn Longstreth tion, and horticulture at the local, state, Susan Krzywicki Milo Baker (Sonoma County): and federal levels. It monitors rare and Administrative Asst: Marcy Millett Lisa Giambastiani endangered plants and habitats; acts to Mojave Desert: Tim Thomas STAFF – AT LARGE save endangered areas through public- Monterey Bay: Brian LeNeve ity, persuasion, and on occasion, legal Fremontia and CNPS Bulletin Editor: Mount Lassen: Catie Bishop Bob Hass action; provides expert testimony to Napa Valley: Gerald Tomboc Legislative Consultant: government bodies; supports the estab- North Coast: Larry Levine Vern Goehring North San Joaquin: Alan Miller lishment of native plant preserves; spon- East Bay Conservation Analyst: Orange County: Nancy Heuler sors workdays to remove invasive plants; Mack Casterman Redbud (Grass Valley/Auburn): and offers a range of educational activi- Development Consultant: ties including speaker programs, field Sandy McCoy Joan Jernegan trips, native plant sales, horticultural Website Coordinator: Mark Naftzger Riverside/San Bernardino: Katie Barrows workshops, and demonstration gardens. PROGRAM ADVISORS Sacramento Valley: Glen Holstein Since its founding in 1965, the tradi- Rare Plant Program Senior Advisor: San Diego: David Varner tional strength of CNPS has been its Jim André San Gabriel Mountains: Orchid Black dedicated volunteers. CNPS activities Vegetation Program Senior Advisor: San Luis Obispo: Kristie Haydu are organized at the local chapter level Todd Keeler-Wolf Sanhedrin (Ukiah): Geri Hulse- where members’ varied interests influ- CNPS Press Director: Stephens ence what is done. Volunteers from the Nancy Morin Santa Clara Valley: Judy Fenerty 33 CNPS chapters annually contribute Poster Program: Bertha McKinley, Santa Cruz County: Deanna Giuliano in excess of 97,000 hours (equivalent Wilma Follett Sequoia (Fresno): Paul Mitchell to 46.5 full-time employees). BOARD OF DIRECTORS Shasta: Ken Kilborn CNPS membership is open to all. Laura Camp (President); David Bigham Sierra Foothills (Tuolome/Calaveras/ Members receive the journal Fremontia (Vice President); Carolyn Longstreth Mariposa): Robert Brown three times a year, the quarterly state- (Secretary); Nancy Morin (Treasurer); South Coast (Palos Verdes): David Berman wide CNPS Bulletin, and newsletters At-Large: Kristie Haydu, Bill Hunt, from their local CNPS chapter. Tahoe: Brett Hall Gordon Leppig, David Varner, Michael Willis L. Jepson (Solano): Vasey, Steve Windhager; Chapter Mary Frances Kelly-Poh Disclaimer: Council Representatives: Orchid Black, Yerba Buena (San Francisco): The views expressed by authors published Glen Holstein Ellen Edelson in this journal do not necessarily reflect established policy or procedure of CNPS, MATERIALS FOR PUBLICATION and their publication in this journal should CNPS members and others are welcome to contribute materials for publication not be interpreted as an organizational in Fremontia. 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FREMONTIA VOL. 42, NO. 1, JANUARY 2014 CONTENTS

EDITORIAL by Kara Moore ...... 2 FLORISTIC DIVERSITY AND DISCOVERY IN THE CALIFORNIA DESERT by James M. André ...... 3 The California desert flora is surprisingly diverse and a hotbed for taxonomic discovery. Looming impacts, however, pose a major threat to this unparalleled floristic frontier.

RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS: PRIORITIES FOR RESEARCH AND CONSERVATION Kara A. Moore and James M. André ...... 9 Many of California’s rare and unique desert plants are at risk, and research on their biology and distribution is urgently needed to fuel effective conservation strategies in our rapidly changing deserts.

APPLYING SPECIES DISTRIBUTION MODELING TO IDENTIFY RARE SPECIES HOT SPOTS by Patrick McIntyre ...... 15 How researchers are using predictive mapping tools to understand plant diversity in California’s deserts.

THE CASTLE MOUNTAINS: A HOTSPOT OF DESERT PLANT DIVERSITY by Duncan Bell ...... 17 Recent explorations of the Castle Mountains in the Eastern Mojave show that they are rich in botanical diversity.

MICROBIOTIC SOIL CRUST COMMUNITIES: A CRITICAL COMPONENT OF CALIFORNIA’S DESERTS by Nicole Pietrasiak and Jeffrey R. Johansen ...... 18 Microbiotic soil crusts are in dire need of conservation because they are essential to desert ecosystems and harbor unexplored biodiversity.

DESERT STREAMS: AN INNOVATIVE NEW APPROACH TO THEIR INVENTORY AND MAPPING by Carolyn Chainey-Davis ...... 20 A new science-based method for mapping California’s episodic desert streams incor- porates an understanding of their unique physical forms and processes where other methods have failed.

THE ECOLOGICAL IMPORTANCE OF WASHES TO DESERT BIRDS by Andrea Jones and Garry George ...... 24 Audubon Society staff share a brief summary of the importance of perennial desert streams and microphyll woodlands to migratory songbirds and other bird species in Southern California.

JOHN O. SAWYER, JR.: 1939–2012 by James P. Smith, Jr...... 25 BOOK REVIEWS ...... 27

THE COVER: Sunrise on the east side of the Granite Mountains in the Mojave National Preserve. In the foreground is a skeleton of buckthorn cholla (Cylindropuntia acanthocarpa) and cinchweed (Pectis papposa var. papposa). September 2013. Photograph courtesy of Stephen Ingram, www.ingramphoto.com.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 1 The first winter storm of the season on Clark Mountain, October 2013. Hidden beyond the mountain lies Ivanpah Solar Electric Generating Facility, covering over 4,000 acres of former desert wilderness. It is expected to power approximately 140,000 homes when it goes on the grid later this year. Photograph by Kara A. Moore. EDITORIAL by Kara A. Moore

his is a momentous time to focus trients cycle. Deserts are rich and di- fornia deserts. Then we focus on the the lens of Fremontia on the verse landscapes in which we can ob- incredible taxonomic diversity of TMojave and Sonoran Deserts. We serve the great drama of nature’s cycles. plants in general and on the biology of teeter on the brink of vast changes in They invite adventurous exploration rare plant species. We also emphasize the North American deserts as we by our city- and computer-trained eyes the impact that periodic water has in expand our cities, roads, recreational as they meet the wild horizon. shaping desert ecosystems. activities, and energy generation into On my first trip across the Mojave The second Mojave and Sonoran one of the largest remaining frontiers and Sonoran Deserts from forested Deserts issue, slated for distribution in on the continent. Policymakers and Michigan, I was struck by the heat and May of 2014, will cover a broad range land managers are embroiled in par- by the expansive landscapes and mes- of management questions. These in- ticular on debate over the siting of merized by the seemingly empty yet clude a strong focus on the ecological utility scale solar and wind renewable colorful mountains, dry lake beds, and consequences of natural and man- energy facilities. sweeping plateaus. Later, when I re- made disturbances. As decisions are Despite the fantastic ability of turned as a researcher exploring on rapidly made regarding resource use, desert ecosystems to withstand drought, foot, I had the gift of time to see closely, we need to challenge each other to endure floods, and capitalize on fa- to witness change over the course of cross institutional boundaries and vorable conditions when they occur, days and seasons. share information including, but not what they cannot recover from—at During my first desert research limited to, species occurrences, popu- least not on the scale of human life project at Ash Meadows National Wild- lation dynamics, and management fail- times—are disturbances so intense and life Refuge, I saw the desert unfold. I ures and successes. To that effect, the sizeable that they interrupt entirely witnessed its dynamic plants and pol- second desert issue will include ar- the flow of soil, water, and species linators, lightening quick leopard liz- ticles from conservation and restora- over the landscape. The next two is- ards and charismatically slow desert tion practitioners and from participants sues of Fremontia focus on the Mojave tortoises, azure pools and pupfish, in policy debates. and Sonoran Deserts and are designed monarch and sphinx caterpillars, rag- There are many additional topics to raise awareness among the public, ing flash floods, and moving soils. On of desert ecology not included in these policymakers, and renewable energy each visit, be it to the lava fields around two issues. Should these issues of proponents of these threats and make Pisgah Crater or the bajadas descend- Fremontia peek your curiosity, Cali- the strong case that deserts are unique ing from Clark Mountain, I continue fornia Deserts: An Ecological Rediscov- and precious ecosystems that warrant to find more diversity, more abundant ery, written by long-time CNPS mem- vigilant protection. life, and more interesting questions ber and contributor Bruce Pavlik, is an The common misconception is that than on the one before. Deserts chal- excellent read for scientists and non- deserts are hostile wastelands, that they lenge us—as they do all of their inhab- scientists alike, and covers topics of are useless open spaces that may be itants—to be patient, to look closely, history, ecology, and management. But carelessly managed. Though they have to build tolerance and resilience. better yet, visit the deserts yourself, intense climates, deserts are anything With this issue of Fremontia and and visit more than once. Try to see but barren. Even in the most extreme the next, we challenge readers to look the deserts in every season. You are places and times, such as those iconic closely at a region that is undergoing likely to gain a new, and deeper appre- and seemingly barren landscapes of radical physical and landscape level ciation for these rich desert ecosys- Death Valley, highly specialized insects change. In this issue, we first highlight tems, and for the great diversity of life and plants persist, soils move, and nu- the diversity and ecology of the Cali- forms that they support.

2 FREMONTIA VOL. 42, NO.1, JANUARY 2014 FLORISTIC DIVERSITY AND DISCOVERY IN THE CALIFORNIA DESERT by James M. André

o other state in the US lore as barren and lifeless places, the and coastal redwood trees. It is also can rival California’s flo- flora of the California desert is in fact where we find California’s shortest- ristic diversity. The Cali- extraordinarily diverse. At present, lived vascular plants, such as the fornia Floristic Province, approximately 2,450 native vascular ephemeral summer annuals that can Nwhich makes up the western two- plant taxa (henceforth termed “spe- germinate and produce viable seed thirds or cismontane portion of the cies” in this article) have been docu- in just three weeks. These same seeds state, is celebrated widely among bi- mented in the California desert, rep- may then lie dormant in the soil for ologists for its unparalleled diversity resenting 38% of the native species many decades before germinating and high degree of endemism, and in the state. If the California desert again. And to underscore the qual- much of California’s extraordinary region was a US state, its flora would ity of the California desert ecosys- diversity is attributed to this unique rank an impressive 18th among states tem, naturalized alien species make region. This review, however, will in total native vascular plant diver- up only 8% of the flora, compared focus on the other floristic provinces sity. Even when compared to the re- to greater than 20% for the rest of of California that comprise the state’s nowned species-rich cismontane the California flora. arid transmontane deserts, a region California, the desert fares quite well. often overshadowed (if not also rain- Mid-elevation alluvial fans of the east- A FLORISTIC FRONTIER shadowed) by the California Floris- ern Mojave Desert support perhaps tic Province, yet of equal significance the highest shrub diversity in all of With more than 100 major and appeal to botanists. California, and similar overall spe- mountain ranges, myriad canyons, The California desert, as defined cies diversity (90–120 total by The Jepson Desert Manual species per hectare) to that of FIGURE 1. THE CALIFORNIA DESERT (Baldwin et al. 2002), comprises the the primeval coastal redwood REGION AND ITS MAJOR COMPONENTS. eastern third of the state and in- forests of northwest Califor- cludes the Great Basin Province east nia. The remarkable richness of the Sierra Nevada crest and the of our desert flora is owed in western components of the Mojave part to its exceptional geologic and Sonoran Deserts (Figure 1, page and topographic diversity, as 3). This vast region is more than well as the rapid speciation twice the size of the Sierra Nevada, and diversification of large spanning 500 miles from the genera (e.g., Phacelia, Astraga- Sweetwater and White Mountains lus, Eriogonum, Cryptantha, of Mono County, southward through Gilia). Death Valley National Park, Mojave Though often described as National Preserve, and Joshua Tree homogeneous, the desert re- National Park, to the lower Colo- gion is in fact a place of great rado Desert of San Diego and Impe- variation and extremes. It is rial Counties. Consisting mostly of where the hottest and coldest federal public lands, largely unpopu- temperatures in the state are lated and still unfragmented by de- recorded, and contains the velopment, the California desert is lowest and nearly the highest of global significance, as it repre- elevations, and certainly the sents perhaps the largest intact eco- driest climate. The desert is system in the US outside of Alaska. where we find the oldest vas- cular plants in California, EXCEPTIONAL DIVERSITY such as creosote bush (Larrea tridentata) and bristlecone Although deserts in general are pine (Pinus longaeva), more Source: Patrick McIntyre, UC Berkeley, 2013, modi- commonly portrayed in literature and ancient than the giant sequoia fied from The Jepson Manual, 2nd edition.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 3 TOP: The Castle Peaks region of the Mojave National Preserve, a rich flora only recently documented by botanists. • ABOVE: Early spring bloom of brittlebush (Encelia farinosa) on the eastern slopes of the unexplored Dead Mountains near Needles, California. All photographs by the author. playas, alkali meadows, badlands completed our floristic inventory of and sprawling sand dune complexes, the California desert, and that the the California desert remains rela- remaining hotbeds for taxonomic tively unexplored. Yet there is a con- discovery are limited to places like tinuing misconception that we have Indonesia and the Brazilian Ama-

4 FREMONTIA VOL. 42, NO.1, JANUARY 2014 zon. This perspective is not entirely mento, and Chemeheuvi Mountains, as Marcus E. Jones, Willis Jepson, surprising given that in the eastern remained virtually uncollected un- Phillip Munz, and Robert Thorne US, and even much of the west in- til very recently. There are no records explored the region extensively. cluding cismontane California, field from the southern half of the Old Though there has been a recent in- exploration and collecting has been Woman Mountains, one of the larg- crease in general collecting by sev- fairly comprehensive over the past est ranges in the eastern Mojave eral botanists, much of our her- two centuries. Desert. Although parts of the Kla- barium collections were made more To gain a better understanding math Mountains, southern Sierra than 50 years ago. of how well we have documented Nevada, and Modoc Plateau are simi- The vast majority of records in the desert flora, I conducted an larly unexplored botanically, no the desert occur along the relatively online search of herbarium voucher other region in the state can match few paved roads in the region. And records, sorted by local physio- the scale of the California desert as a while the more inaccessible areas graphic areas (mountain ranges, floristic frontier in need of docu- remain major spatial voids for docu- valley basins, etc.). To date, most mentation. mentation, temporal voids exist as desert ranges and basins have fewer Several other important obser- well. Summer annuals and fall-flow- than 250 herbarium records in the vations stand out when viewing the ering plants are prominent in the Consortium of California Herbaria. collections history for the Califor- California desert, comprising about And some ranges such as the Dead, nia desert. Efforts to inventory and 40% of the flora, but poorly docu- Hackberry, Resting Spring, Ship, document desert plants, as measured mented because a disproportionate Owlshead, and West Riverside by the number of vouchers collected number of collections have been Mountains, have fewer than 100 per decade, have actually declined made in spring and early summer. records. Several major ranges, in- since the first two-thirds of the 20th Major information gaps still exist, cluding the Turtle, Castle, Sacra- century when famous botanists such and I make the case here that we

Sunrise on the impressive volcanic spires of the northern Turtle Mountains, a region unexplored by botanists until this past decade.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 5 Late summer bloom of cinchweed (Pectis papposa) in the Granite Mountains of eastern San Bernardino County, showcasing the often- overlooked floristic diversity of autumn in the desert. have only scratched the surface in the San Diego County Plant Atlas Until recently, the Whipple our understanding of the desert flora. Project (desert portion) that began Mountains of southeastern San Ber- What can we expect to add to the in 2003 (Rebman et al.), a flora of nardino County represented one of California desert flora with further Joshua Tree National Park (La Doux), the many botanical black holes in exploration? Recent floristic studies a flora of Mojave National Preserve the California desert. Sarah De Groot demonstrate what can be accom- and surroundings (André), and flora collected the 500-square-mile range plished with robust and focused in- projects by botanists from Rancho extensively in the mid-2000s as part ventory and assessment. These ef- Santa Ana Botanic Garden (e.g, of her graduate research, and docu- forts include, but are not limited to Honer, Fraga, Bell, and De Groot). mented approximately 400 species,

6 FREMONTIA VOL. 42, NO.1, JANUARY 2014 a four-fold expansion of the flora TOP TO BOTTOM: Examples of desert species (De Groot 2007). In the process of new to science. A close-up of the unique flower of the rare shrub Orocopia Moun- adding many common species, she tains spurge (Euphorbia jaegeri). • The discovered several new species in showy Clark Mountain monardella (Mon- the state and the California desert ardella eremicola), one of several new flora, and documented numerous species of this genus that have recently been added to the desert flora. • The showy significant range extensions. Clark Mountain green-gentian (Frasera In 2003 Joshua Tree National albomarginata var. induta), only found in Park (JTNP) launched a compre- California on the limestone substrates of hensive effort to inventory the park’s Clark Mountain. 1,200 square miles. Prior to this ef- fort, many areas in the park such as Cinder Cones, Castle Peaks, Piute the Pinto Basin, Long Canyon, and Range, and Woods Mountains, the Coxcomb and Eagle Mountains where few to no historic vouchers remained virtually unexplored. Led existed. Over the course of the by park botanist Tasha La Doux, the project I focused approximately goal of this project was to build upon 9,000 field hours upon these voids, previous efforts, with an emphasis and also continued to survey the on surveying the under-explored higher ranges and Kelso Dunes spatial and temporal gaps. Over the (Thorne et al. 1981). Since that time, past 10 years the inventory at JTNP 120 native species have been added has added 140 species to the park to the flora (a 14% increase), many list, expanding the list by 24% to its of which were found in the “well- current 725 species. Roughly 90% studied” areas (e.g., New York and of the additions were made via field Providence Mountains). Seven of discoveries, including five new spe- these species are new to science, cies to science, while 10% were more than 800 new rare plant oc- added through herbarium searches currences have been added, and and taxonomic revisions. noteworthy range extensions con- The final case study presented tinue to be frequently documented. here is my own work on a flora of Assuming the three flora projects the Mojave National Preserve (MNP) outlined above are representative of and surrounding areas (>3,000 the undocumented status of the en- square miles), initiated in 1995. Un- tire California desert, we can take like most of the California desert, home one clear message: the desert the MNP region received consider- remains a floristic frontier, and these able attention from early botanists. are not the end of days for taxo- In particular, the Clark, New York, nomic discovery. On the contrary, I Providence, and Granite Moun- would suggest that we are in the midst tains—the highest ranges in the Pre- of a second golden age of discovery. serve—have attracted a number of prominent collectors over the past 150 years. Robert Thorne, Barry RECENT DISCOVERIES AND Prigge, and James Henrickson ex- ADDITIONS plored parts of the MNP extensively about 35–40 years ago, and pub- To estimate the rate of additions lished a flora that included the Kelso of native plant species to the Cali- Dunes and the high ranges. fornia desert, I utilized the popular In 1995 I mapped the approxi- floras that have been published over mately 15,000 historic vouchers the past 90 years. The numbers of known from the region at the time desert species included in each pub- and noticed that nearly all were clus- lished flora are as follows: tered in the Kelso Dunes and four high ranges. There were 16 other 1925 Jepson CA Flora: 2,149 mountain ranges, such as the Cima 1973 Munz & Supplement: 2,187

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 7 1993 Jepson Manual: 2,267 190–200 native species to be added places on earth, this is your call to 2012 Jepson Manual, 2nd ed: 2,430 to the California desert flora by the action. 2013 (Present): 2,450 year 2100. This projection assumes we have reached the peak rate of REFERENCES While there has been an increase discovery (33 species per decade), of approximately 300 species over and applies a 10% decline in the André, J.M. 2006. A vascular flora of 90 years, representing a 14% in- number of additions per decade go- the Granite Mountains, Eastern San crease, more than 60% of these were ing forward. Another way of look- Bernardino County: An annotated added in just the last two decades. ing at this projection is to consider checklist. Crossosoma 32(2): 38–74. The lull in the mid-1900s coincided what it means to our present level André, J.M. 2005. Inventory of Vascu- lar Plants at Mojave National Pre- with a national lumping trend in of understanding. If 200 species are serve: A Preliminary Report. Tech- plant taxonomy that occurred dur- to be added to the California desert nical Report #2280201178, Inven- ing that period. Most of the addi- by the end of this century, and per- tory & Monitoring Program, US tions are newly described species, haps another 50–100 more in the Department of the Interior, National but some represent species previ- next century, then we can safely Park Service, Barstow, CA. ously known only from adjacent assume that at least 10% of the flora Baldwin, B.G., et al., eds. 2012. The states or bioregions of California. In is presently undescribed. This is Jepson Manual: Vascular Plants of the past two decades the desert flora quite humbling. California. UC Press, Berkeley, CA. has expanded by 183 species or 8%. The projection of 200 new addi- Baldwin, B.G, et al., eds. 2002. The Over the same period, the Califor- tions this century also assumes that Jepson Desert Manual. UC Press, Ber- nia flora (including the desert) has the concept of a species remains rela- keley, California. Barbour, M.G., and J. Major. 1988. Ter- grown by only 5%, suggesting that tively unchanged and that extinc- restrial Vegetation of California. Cali- the California desert is indeed a ma- tion is not a factor (i.e., that the fornia Native Plant Society, Davis, jor hotbed for taxonomic discov- desert will remain sufficiently pro- CA. ery. In addition, I am aware of at tected into the future). As to the De Groot, S.J. 2007. The “nose” of Cali- least 40 proposed new desert spe- latter point, approximately 1,500 fornia: An important part of the cies likely to be added to the flora square miles of the California desert state’s diversity. Fremontia Vol. 35 this decade. are currently proposed for indus- (No 1): 3–6 This resurgent golden age of dis- trial-scale solar and wind energy de- Hickman, J.C. ed. 1993. The Jepson covery is by no means as high as the velopment in the next several years, Manual of Higher Plants of Califor- late 1700s and 1800s when the first mostly on undisturbed federal lands. nia. UC Press, Berkeley, CA. wave of exploration occurred, but Considering indirect impacts of de- Jepson Flora Project. 2013. Jepson eFlora, http://ucjeps.berkeley.edu. it’s happening now during a time velopment (e.g., increased vectors Jepson, W.L. 1925. A Manual of the when field collecting (and funding for alien species invasions, losses to Flowering Plants of California. UC for taxonomy) has seen a general biological soil crusts, barriers to dis- Press. Berkeley, CA. decline. The current taxonomic ten- persal, etc.), the actual degradation McLaughlin, S.P. 1995. Floristic rela- dency to split species, combined with of habitat will extend far beyond the tionships of the eastern Mojave the fact that more people are now footprint of the projects themselves. Desert: A quantitative analysis of lo- describing new species than had been According to most extinction mod- cal floras. Crossosoma 21:57–74. the case 50 years ago (and doing so els, at this unprecedented scale and Munz, P.A. 1959. A Flora of California. more sensibly I believe) explains pace of impact, extinction events will UC Press, Berkeley, CA. some of this recent increase in num- be likely. Stein, B.A. 2002. States of the Union: bers of species in the California My hope is that we continue Ranking America’s Biodiversity. A Natureserve Report. The Nature desert. Tools such as Google Earth this push to inventory the Califor- Conservancy, Arlington, VA. and the availability of online data- nia desert, as we still have much Thorne, R.F., B.A. Prigge, and J. bases such as that of the Consortium work to do, and time is now of the Henrickson. 1981. A flora of the of California Herbaria also greatly essence. Perhaps more important is higher ranges and the Kelso dunes improve the efficiency of fieldwork, that the desert gains the level of of the Eastern Mojave Desert in Cali- especially when targeting remote ar- appreciation, advocacy, and protec- fornia. Aliso 10(1): 71–186. eas that are difficult to access. tion that it deserves. To both grizzled and nascent students of botany, the James M. André, University of California WHAT THE FUTURE HOLDS California desert holds a wealth of Riverside, Department of Biology, UCNRS research and teaching opportuni- Granite Mountains Desert Research Based upon fairly conservative ties. And to those interested in pre- Center, HC1 Box 101, Kelso, CA 92309, assumptions, we can expect another serving one of our last big wild [email protected]

8 FREMONTIA VOL. 42, NO.1, JANUARY 2014 California bearpoppy (Arctomecon merriamii), a rare perennial found in small populations on calcareous gravel deposits of the northeastern Mojave Desert and in western Nevada. Photograph by Erica Smith. RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS: PRIORITIES FOR RESEARCH AND CONSERVATION Kara A. Moore and James M. André

mong people of all cultures function and the role that each life in our deserts because they have there is an inherent ap- form plays is incomplete. The unique systematically received far less at- preciation of things that and diverse array of rare plants in tention from botanists than other are rare and unique, the California deserts presents such bioregions. In the California deserts, whetherA a precious gemstone or a conservation challenge because the native plants are being added to the green flash on the horizon at sunset. distributions and basic life histories flora faster than any other region of To the conservation scientist, rare of most species remain poorly docu- the state. For more on this excep- organisms are of particular interest mented. tional floristic frontier, see the other because they are genetically distinct While endangered mammals and article by James M. André on page 3 aspects of living diversity that may birds often command more public of this issue. be at risk of extinction, or may sig- attention, plants form the basis of The California Native Plant nify an impaired ecosystem. As stew- all terrestrial life by providing en- Society’s (CNPS) Inventory of Rare ards of the planet, most of us believe ergy, materials, and habitat struc- and Endangered Plants ranks 35% that we have an ethical obligation to ture. Globally, 25–40% of native vas- of California’s native vascular flora preserve biodiversity and the eco- cular plant taxa (henceforth termed as being of conservation concern, logical processes that sustain natu- “species” in this article) are at risk but only 20% of the California desert ral systems. This goal is especially (Pitman 2002). Within North flora is similarly ranked. There are challenging, given that our under- America, numbers of imperiled at least three reasons for this lower standing of how healthy ecosystems plants may be most underestimated percentage. First, endemism (where

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 9 Fantastic bloom in the foreground of the rare Bartow wooly sunflower (Eriophyllum mohavese)—a plant that is often less than a centimeter in width and height—in 2011 near Kramer Junction in the northwestern Mojave Desert of California. In infrequent favorable years, these tiny rare sunflowers form an unusually dense carpet under the common annual Bigelow’s tickseed (Coreopsis bigelovii). Photograph by Kara Moore. Close-up photograph by James André. a species is only found in a restricted ticularly given that the region is a basis for establishing strategies for habitat or location) is lower in the hotbed for new species discovery— conservation. Our goal is to bring California deserts than in the rest of so most new additions to the flora attention to the diversity of rare California. Yet even though the Cali- will be rare. desert plants and to provide insight fornia deserts represent only a frac- The California deserts are un- on their ecology that will be useful tion of the Great Basin, Mojave, and dergoing rapid change as a result of to those planning for their protec- Sonoran Deserts, a striking 25% of increasing use by humans, especially tion. We use case studies to illus- the rare species in the California on federal public lands targeted for trate some of the unique facets of deserts are endemic to the state, widespread solar and wind energy rare desert plants that influence their highlighting the uniqueness of the development. Planning measures for management, highlighting addi- California desert flora within the rare plant protection will be based tional research needs and priorities. North American deserts. Second, upon limited existing knowledge of because the deserts are, at present, the desert flora, and thus lack the relatively pristine compared to much necessary level of detail. Even after OPPORTUNISTS BY of California, most rare desert plants we document the distributions of NECESSITY are inherently rare, with only a hand- rare plants, we are challenged to ful driven to rarity by human activi- understand their unique biology. Desert plants must be adapted ties. Third, we have underestimated In this article we focus on key to a harsh and variable climate to the actual number of rare plant spe- factors that influence rarity in the persist. These adaptations include cies in the California deserts—par- California desert flora and provide a the climatic responsiveness of an-

10 FREMONTIA VOL. 42, NO.1, JANUARY 2014 nuals, the seasonality of herbaceous thetic pathway (C4) than and woody perennials, and the en- the majority of plant spe- during architecture of many succu- cies (C3), which allows lents and shrubs. Desert annuals them to more efficiently arise from a long-lived dormant seed grow and reproduce in this bank only when triggered by the warm climate with sparse right combination of temperature and/or unpredictable rain- and moisture. Seed dormancy buff- fall. ers annual populations from climatic The challenge of limited fluctuations by permitting emer- field documentation and gence only when conditions are periodic emergence is ex- likely to lead to high survival (Clauss emplified by Mojave milk- and Venable 2000). Many desert weed (Asclepias nycta- annuals have a very limited period ginifolia), a tuber-forming during which germination and emer- perennial limited to the gence can occur, and it is highly eastern Mojave. Prior to sur- dependent upon precipitation. In veys for the construction of some cases, especially in the sum- the Ivanpah Solar Elec- mer, this window may be only a few tric Generating System weeks. The absence of aboveground (ISEGS)—a massive project plants over multiple years can cre- that has been built on ap- ate the false impression that plant proximately 3,740 acres of populations are no longer present, federal public land—the although they are merely lying in milkweed was known in wait as viable seed in the soil for California from only four favorable germination conditions. occurrences, all above 3,000 One plant that demonstrates this dra- feet in elevation, and all at matic year-to-year variation is the some distance from the miniscule Barstow wooly sunflower project site. Mojave milk- (Eriophyllum mohavense), a rare win- weed was thus assumed to ter annual found only near Kramer have low potential to occur Junction and Edwards Air Force Base in Ivanpah Valley and was (photo on page 10). overlooked during initial project sur- The long dormancy periods of veys, which were conducted during rare herbaceous desert perennials fairly dry years. Follow-up surveys must be considered in the assess- in subsequent years of increased ment of population densities and in rainfall revealed numerous popula- developing long-term monitoring tions in the ISEGS footprint. These programs. The flowering phenology plants had been dormant and unac- of some herbaceous perennials can counted for during the preliminary be highly complex. For example, environmental review process. scarlet four o’clock (Mirabilis cocci- nea) typically flowers in response to RARE PLANT HABITATS localized summer precipitation, but may also respond to late winter rains The causes of rarity in the desert and then flower in early summer. are nearly as diverse as the rare flora Some deciduous shrubs such as vera itself (e.g. Rabinowitz 1981; Fielder dulce (Aloysia wrightii) and desert 1986, 2001). Many rare desert spe- Desert milkwort (Polygala acanthoclada) milkwort (Polygala acanthoclada) cies are narrow endemics that occur in bloom. Photographs by James André. remain leafless until they receive in a single restricted geographic area, sufficient precipitation. Many mon- and/or may be confined to a unique that have become restricted and now soon-responsive summer- and fall- habitat to which they have adapted. persist in an isolated location such flowering desert species such as They may have either recently as a mountaintop. Unique desert desert purslane (Portulaca hali- evolved in place (neoendemics), or habitats rich in endemic rare spe- moides) apply a different photosyn- be ancient species (paleoendemics) cies include characteristic uplands

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 11 such as sand dunes, gypsum clay habitats, their distribution and rar- fluence rarity, the production and deposits, and limestone substrates. ity being driven by other factors. fates of seeds are often the most Deserts also include localized wet- These include a long list of taxa critical and least understood, and land ecosystems that are home to such as Orocopia Mountains spurge the complex interaction of flowers many rare plants, including alkali (Euphorbia jaegeri), Mojave mon- and fruits with animals warrants ex- meadows, non-alkali wetlands, keyflower (Mimulus mohavensis), tensive research. Some rare species freshwater springs, riparian systems, and the crucifixion thorn (Castela employ self-incompatibility, mean- and infrequently inundated land- emoryi). These are fragmented or ing that individuals need pollen from scape features such as playas and disjunct populations that are diffi- another individual to produce vi- washes (Pavlik 2008). cult to explain ecologically. Causes able seed. These species, such as Another challenge for rare habi- of rarity for such species might in- Little San Bernardino Mountains tat specialists is the limited extent, clude geographic isolation due to linanthus (Linanthus maculatus), fragmentation, and disjunction of long-term climate change, altered need larger population sizes to per- many of their specialized habitats. pollinator guilds, restricted gene sist than species that readily self- For example, species specialized on flow, or other factors. Additional pollinate. gypsum clay (gypsophiles) and lime- research is often required to discern Idiosyncrasies of rare plants in- stone substrates such as California whether such widely scattered popu- clude atypical chromosome num- bearpoppy (Arctomecon merriamii) lations are genetically isolated across bers, low genetic diversity, and rela- have populations scattered through- the range of the species’ distribu- tionships with specialist pollinators out suitable habitat patches that can tion. or dispersers. And these factors com- be separated by many miles. Lime- monly compound to influence rare stone ranges that extend into Cali- species. For example, Peirson’s milk- fornia from Nevada, such as Clark INFLUENCES OF vetch (Astragalus magdalenae var. Mountain and the Nopah Range, REPRODUCTIVE BIOLOGY peirsonii) is restricted to the sands house a suite of limestone special- of the Algodones Dunes, is self-in- ists that are rare in California, but In developing conservation strat- compatible, and requires pollination fairly common in Nevada where egies for rare desert plants it is of cri- by a single native bee species (Groom limestone substrates are abundant. tical importance to first understand et al. 2007). In addition, its habitat Rare species may also be distrib- their diverse reproductive systems. has been severely degraded by off- uted across a range of ubiquitous Of the many factors that can in- road-vehicle use. Even when seeds are produced, their ability to sur- vive, disperse, and persist in the seed bank is not assured. Rare plants at A PROFILE OF CALIFORNIA’S RARE DESERT FLORA Ash Meadows and Pisgah Crater can lose up to 90% of their total seed he 491 listed rare California desert species are an exceptionally production due to herbivory by jack- Tdiverse group with representatives in 70 vascular plant families. rabbits, sometimes for several con- Nearly a quarter of these species (152) are either endemic to the secutive years (Pavlik et al. 2009). California deserts or rare both here and elsewhere, placing them in Rapid construction of large-scale the highest rarity class, California Rare Plant Rank (CRPR) 1B. A energy projects in the California large proportion (209 species, 43%) are CRPR 2B, meaning they are desert could exacerbate seed losses rare in California but more common in other states. This is a rela- due to local distortion of food webs. tively high percentage of 2B species in comparison to the rest of California. Another 118 plants are listed as CRPR 4, a watch list of species with limited distributions. NEEDS AND PRIORITIES Of the CNPS-listed desert species, only 13 have federal listing We must put our feet to the earth. status (are protected) under the Endangered Species Act, with 8 There is an ongoing need for inven- receiving protection as “Endangered” and the other 5 listed as “Threat- tory and mapping of known rare ened.” Federal listing of desert plants is often sidelined by other plant species in the California deserts critical research needs or because officials may assume that rare to better inform regional conserva- species occurring on federal lands are sufficiently protected. Yet we tion and management. The majority have identified an additional 20–30 desert species that potentially of rare plant records occur within a meet criteria for federal listing status, and listing would greatly mile of major roads, in historically improve their prospects for research funding and protection. popular botanical areas (e.g., New York Mountains, Mecca Hills), or

12 FREMONTIA VOL. 42, NO.1, JANUARY 2014 White-margined beardtongue (Penstemon albomarginatus) in bloom near the Sleeping Beauty Mountains, Central Mojave Desert, California. Despite the numerous flowers in 2011, extremely few fruits and seeds were successfully produced because of strong herbivory and other unknown limits on fruit development. Photograph by Kara Moore. in areas surveyed prior to develop- such as from urbanization, utility- sistence of species. A comprehen- ment. Geographic models can be scale energy, mining, recreational sive life history approach that in- used in some cases to estimate spe- activities, invasive species, and cli- cludes monitoring of all life stages, cies ranges, but have important limi- mate change. At present, status re- including seed banks, is necessary in tations when applied to rare and ports that document basic biology order to make realistic predictions under-surveyed species (see article and perceived threats are available on population viability. Each stage by Patrick McIntyre on page 15). for only about a quarter of the 491 of growth and reproduction can be Given that the California deserts known rare desert plants, and fewer greatly affected by other species in remain a floristic frontier (we have than 5% have accompanying conser- the community, some negatively added 183 species in the past two vation management plans. Of equal (competition and herbivory) and decades alone), extensive inventory concern is the fact that established some positively (pollination, seed is still needed to catalog additions to long-term monitoring or research dispersal). Therefore it is necessary the native flora. Many undiscovered programs exist for less than 1% of all to implement research that includes species will also be rare, and not rare plants in the California desert. interacting species and is conducted afforded the necessary protections We need to increase and extend spe- over several decades. Such a time until their taxonomy is confirmed cies-specific research to provide a frame can better capture population and their distributions mapped. A baseline understanding of a repre- responses to fluctuations in climate comprehensive desert-wide inven- sentative set of rare desert species. and variations in species interactions. tory will take institutional will, much Basic population studies that fol- Unfortunately, such long-term stud- expertise, and many decades to com- low the fates of individual plants ies are extremely rare. plete. and their progeny are especially Currently, we are assembling 13 In addition, research on the ecol- needed to make predictions about years of field data and observations ogy of California’s rare desert spe- the effects of landscape fragmenta- on white-margined beardtongue cies is vital for stewarding popula- tion, population reduction, and cli- (Penstemon albomarginatus), an her- tions amid rapidly increasing threats, mate change on the long-term per- baceous perennial known in Cali-

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 13 fornia from a single population near chortus striatus) and crowned muilla ecosystem and landscape units. Pres- Pisgah Crater. We are assessing how (Muilla coronata) can be nearly im- ervation of large contiguous and di- different life stages, precipitation, possible to observe before herbivores verse wildlands throughout the Cali- and species interactions affect its consume aboveground leaves and fornia deserts is the only way to ability to persist. We found that stems. However, for the majority of ensure protection of its many rare caging plants from black-tailed rare species, by far the most critical species. jackrabbit herbivory increases sur- interactions are those they have di- vival and seed production, but per- rectly and indirectly with humans. REFERENCES haps not enough to protect the spe- cies from other factors that threaten A CALL FOR RESEARCH Clauss, M.J., and D.L. Venable. 2000. local extinction. Increased drought Seed germination in desert annuals: frequency, as predicted by global cli- Ongoing research is needed to An empirical test of adaptive bet mate change models, radically in- determine the diversity, abundance, hedging. American Naturalist 155: creases the probability of extinction. and distributions of rare desert plants 168–186. Fielder, P.L. 1986. Concepts of rarity Additional demographic monitoring and to understand the fundamen- in vascular plant species, with spe- is needed to determine if this popu- tals of their ecology. We emphasize cial reference to the genus Calo- lation will rebound or rapidly ad- that the most critical facet of inven- chortus Pursh (Liliaceae). Taxon 35: vance to California Rare Plant Rank tory and mapping for rare species is 502–518. 1A status (plants presumed extinct to remember that absence cannot be Fielder, P.L. CNPS Inventory, 6th Edi- in the state). assumed from a single survey (or tion, 2001. For many desert rare plants, in- even a few) for species that have a Groom, J.D., et al. 2007. Quantifying cluding the white-margined beard- dormant life phase. And with rapid off-highway vehicle impacts on den- tongue, perhaps the most curious and extensive changes in arid land sity and survival of a threatened and important life stage is the most use, we need to advance taxonomic dune-endemic plant. Biological Con- challenging to study: the soil seed research on both known species and servation 135(1): 119–134. IUCN 2013. The IUCN Red List of bank. Unknown factors include the those that are not yet described in Threatened Species. Version 2013.1. longevity of seeds under field con- order to forward their protection. http://www.iucnredlist.org. ditions, mechanisms and rates of dis- There is a pressing need for popu- Pavlik, B.M. 2008. The California persal, local density, and conditions lation biology research on rare plant Deserts: An Ecological Rediscovery. that trigger germination. Seedling species. Focal points must include University of California Press, Ber- emergence, determined by environ- research on the detection and func- keley, CA. mental factors controlling germina- tion of dormant life stages and the Pavlik, B.M., Moore-O’Leary, K.A., and tion and mortality in the soil, is of- positive and negative effects of spe- A.E. Stanton. 2009. Quantifying her- ten the most critical driver of popu- cies interactions. We must deepen bivore impacts on rare plants at Ash lation growth. When mathematical our understanding of how both dor- Meadows. Proceedings of the Califor- models are used to simulate popula- mant life stages and interactions with nia Native Plant Society Conservation Conference. tions of white-margined beard- other species vary with fluctuations Pitman, C.A., and P.M. Jørgensen. tongue, small variations in the mea- in climate that are characteristic of 2002. Estimating the size of the sured rates of seedling emergence these ecosystems. Furthermore, re- world’s threatened flora. Science (1 can result in very different proba- search to explore the effects of prox- November 2002): 989. bilities of population growth and imity to development, landscape Rabinowitz, D. 1981.Seven forms of persistence. fragmentation, soil erosion, altered rarity. In The Biological Aspects of We also must improve our un- hydrology, and other anthropogenic Rare Plant Conservation, ed. H. derstanding of rare species’ interac- disturbances on rare plants is criti- Synge. John Wiley & Sons Ltd., New tions with other plants, pollinators, cal to their persistence. York, NY. mutualists, and herbivores. Many Regional and global processes plants endure strong herbivory, at such as atmospheric nitrogen depo- Kara A. Moore, Department of Evolution least periodically, when populations sition and global climate change have and Ecology, University of California, of insects, jackrabbits, pocket go- far-reaching effects on local popula- One Shields Avenue, Davis, CA 95616, phers, wood rats, or other species tions and must be included in the [email protected]; James M. reach high densities (often cyclic or scope of studies. Although species- André, Department of Biology, University due to a lack of predators), or dur- specific management is required by of California, Riverside, and UC Natural ing drought when food resources the biological uniqueness of rare spe- Reserve System, Granite Mountains Desert are limited. For example, in some cies, clearly all forms of life are best Research Center, HC1 Box 101, Kelso, years, alkali mariposa lily (Calo- served by protecting functioning CA 92309, [email protected]

14 FREMONTIA VOL. 42, NO.1, JANUARY 2014 APPLYING SPECIES DISTRIBUTION MODELING TO IDENTIFY RARE SPECIES HOT SPOTS by Patrick McIntyre

n the last 15 years, ecologists have then make a map of predicted habi- developed mapping tools to pre- tat for a species beyond areas where dict new areas where rare species it is currently known by identifying may occur, based on their known places with similar habitat. Com- Ilocalities. These tools are broadly bining these tools with predicted referred to as species distribution or changes in temperature and rainfall, ecological niche models. (Ecologists ecologists can also make predictions debate whether the method predicts about where species have the poten- the ecological niche of a species or tial to occur in the future, given dif- Two California endemics known from the merely where the species occurs on ferent scenarios of climate change. Ord Mountains southeast of Barstow, a the landscape.) Species distribution predicted hotspot of rare plant diversity. LEFT: Mohave monkeyflower (Mimulus modeling identifies aspects of the mohavensis). • RIGHT: Creamy blazing star environment, such as elevation, tem- A MAP OF RARE PLANT (Mentzelia tridentata). Both photographs perature, and rainfall associated with HOTSPOTS by the author. where a species is known to occur. Using mathematical models and To illustrate how these tools can rare plant species whose docu- Geographic Information System inform plant conservation, I built mented occurences allowed us to (GIS) technology, ecologists can species distribution models for 151 identify at least 5 populations. Using

PREDICTIVE MAPS OF RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS.

LEFT: MAP OF CRPR 1 PLANTS (PRIMARILY CALIFORNIA ENDEMICS). • RIGHT: MAP OF RARE OR THREATENED PLANTS IN CALIFORNIA AND WHICH OCCUR MORE COMMONLY OUTSIDE OF THE STATE.

Note: Areas in warmer colors (yellows and browns) are areas of higher predicted rare plant diversity. The predicted region corresponds to the proposed boundaries of the Desert Regional Energy Conservation Plan.

Source: Patrick McIntyre, UC Berkeley. Predictive mapping of rare desert plants, 2013.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 15 (Menodora spinescens var. moha- vensis), and is predicted to have suit- able conditions for several other rare species. Based on the simple maps presented here, this area might sup- port additional rare species not cur- rently documented in those loca- tions, or might be a region that could act as a refuge for plants from other areas forced to shift their ranges as a result of climate change. Predictive maps such as the ones accompanying this article represent valuable tools that can help guide field-based efforts to document plant diversity. They can identify unan- ticipated locations where rare spe- cies might be found. They can also be used to help predict where ap- propriate habitat for rare plants could occur over large areas that are infeasible to survey by foot. In addi- tion, they provide a means for pre- dicting where species might occur in the future under scenarios of cli- Four rare plants known from the Clark and Kingston ranges in the eastern Mojave desert mate change, something that can- of California, a known and modeled hotspot of rare plant diversity in California. CLOCKWISE not be accomplished through field FROM TOP LEFT: Stephen’s beardtongue (Penstemon stephensii). Photograph by Jim André. • Mohave milkweed (Asclepias nyctaginifolia). Photograph by Drew Maraglia. • Parish’s club surveys. cholla (Grusonia parishii). Photograph by Kara Moore. • Mojave desert plum (Prunus However, predictive maps are eremophila). Photograph by Jim André. based on imperfect data—known lo- calities that represent only a subset a standard set of climate data for the species from other habitats that ex- of each species’ real distribution. 20th century (worldclim.org), I tend to the edge of California’s They are not substitutes for on-the- made maps of where these plants Mojave and Sonoran Deserts. Sec- ground exploration by experienced were predicted to occur across the ondly, the Clark and Kingston botanists. It’s one thing to use a spe- Desert Renewable Energy Conser- mountain ranges in eastern Califor- cies distribution model to identify vation Plan (DRECP) area. This area nia, much of which are included in likely areas to hunt for new rare is the recent focus of rapid develop- the Mojave National Preserve, are plant occurrences, and another thing ment and conservation planning. I identified as centers of diversity both entirely to use the predictions of a then overlaid predictions for Cali- for endemic and non-endemic taxa. model to decide which parcels of fornia Rare Plant Rank (CRPR) 1 The Preserve’s ranges host many spe- land to preserve and which to de- species (those primarily endemic cies that normally occur outside of velop. Finally, models can’t predict to California) and CRPR 2 species California, but also occur in these the distribution of a species that has (those rare in California but more mountains. never been described, and new spe- common elsewhere) in order to iden- Another intriguing area of pre- cies are described every year in the tify predicted regions of rare plant dicted high diversity is the Ord and California desert. These models are diversity in the California deserts Lavabed mountain ranges southeast useful tools that we can use to guide (see maps on page 15). of Barstow. This area falls outside research and focus field exploration Several patterns emerge from ex- of National Park/National Preserve based on what we know today, and amination of the maps of predicted areas and currently is known to what might likely occur in the fu- diversity. First, western areas of the support a number of rare Califor- ture. desert abutting mountain ranges are nia endemics, such as the creamy identified as areas of high diversity. blazing star (Mentzelia tridentata), Patrick McIntyre, University of Califor- This highlights how these areas sup- Mohave monkey flower (Mimulus nia, 3101 Valley Life Sciences, Berkeley, port both true desert species and mohavensis), and Mojave menodora CA 94720, [email protected]

16 FREMONTIA VOL. 42, NO.1, JANUARY 2014 Joshua tree woodlands amidst a unique assemblage of grasses at sunset in a valley surrounding the Castle Mountains. Photograph by the author. THE CASTLE MOUNTAINS: A HOTSPOT OF DESERT PLANT DIVERSITY by Duncan Bell

he Castle Mountains are a center of the Castle Mountains is The botanical frontier of the fantastic example of the great rugged and rocky, with hidden can- Castle Mountains has only recently diversity of relatively small yons containing a diversity of rare been breached by botanists such mountain ranges in the Cali- desert annuals, including nine- as James André, Andy Sanders, and forniaT deserts. In San Bernardino awned pappus grass (Enneapogon myself. Every year we are finding County, the Castle Mountains are desvauxii) and Clark Mountain new and previously unknown rare only approximately 30 square miles, spurge (Euphorbia exstipulata). plant populations, and certainly yet they are home to over 30 rare Steep canyons spill out into wide more discoveries remain. For ex- plant species and hundreds of com- valleys that surround the range. They ample, in 2012 both James André mon species. Enter any canyon of are home to dense and extraordinar- and I found Mexican panicgrass the Castle Mountains, scramble onto ily healthy stands of Joshua trees that (Panicum hirticaule) in the Castle any ridgeline, or walk any wash and are part of a desert savannah con- Mountains area, the first collections you will find unique and interesting taining a diversity of native annual of this species for San Bernardino plant species, some of which are and perennial grasses. Around two County and an 88-mile northern found only in this rugged corner of dozen grass species are found here, range extension. the Mojave, such as canyon bird’s of which half a dozen are rare grasses. Every field press that is filled foot (Lotus argyraeus var. multi- Some species, such as burro grass with care in the Castle Mountains is caulis) and the showy pinto beard- (Scleropogon brevifolius) and false likely to hold important discoveries tongue (Penstemon bicolor). buffalo grass (Munroa squarrosa), are that will allow us to further under- The Castle Mountains, publicly part of unique desert grasslands stand our rare plant populations and owned by the Bureau of Land Man- found nowhere else in California. their distribution in the rugged but agement, are best accessed by Hart Also present are many other rare plant fragile California deserts. Mine Road from the Mojave National populations such as matted cholla Preserve. Access requires a high (Grusonia parishii), Abert’s sanvitalia Duncan S. Bell, 1500 North College Ave- clearance vehicle, and four-wheel aster (Sanvitalia abertii), and red four nue, Claremont, CA 91711-3157, dbell drive is seasonally necessary. The o’clock (Mirabilis coccinea). @rsabg.org

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 17 MICROBIOTIC SOIL CRUST COMMUNITIES: A CRITICAL COMPONENT OF CALIFORNIA’S DESERTS by Nicole Pietrasiak and Jeffrey R. Johansen

ow more than ever, pris- are especially crucial to the ecologi- of converting atmospheric nitrogen tine and fragile desert cal functioning of desert ecosystems. to ammonium, an essential but lim- ecosystems are facing The crusts harbor diverse taxa in- ited nutrient in desert systems. Thus, major threats from hu- cluding mosses, lichens, fungi, green crusts represent an important nitro- Nman development, and efforts are algae, diatoms, and cyanobacteria gen source for associated vascular being made to protect sensitive, spe- that bind together mineral soil par- plant communities or soil food webs. cial-status plant and animal species. ticles into water- and wind-stable Biological soil crusts also can seques- In contrast, microbial communities aggregates at the soil surface. This ter substantial amounts of carbon. of desert soils have been neglected crust is vital because it prevents ero- The carbon can accumulate in the in conservation efforts, although they sion in sparsely vegetated landscapes. microbial biomass, ultimately add- are often considered as critical to In addition to their important ing soil organic matter to the sys- ecosystem health. It is well docu- role in stabilizing soil surfaces, their tem. This organic matter is then avail- mented that the microbial commu- contribution to soil fertility is essen- able in the soil food web. In fact, nities within biological soil crusts tial. Some crust microbes are capable biological soil crusts are rightfully

Biological soil crusts of Californian Deserts. • BELOW: The darkened soil surface indicates the presence of lichen and algal crusts. The measuring tape gives an idea of scale. • FACING PAGE, TOP: An algal crust is all that holds together soil particles on this desert slope, which contains little vegetation. • FACING PAGE, BOTTOM: A picture taken with a microscope of a potential new cyanobacterial genus and species found in a wilderness area of Joshua Tree National Park. All photographs by Nicole Pietrasiak.

18 FREMONTIA VOL. 42, NO.1, JANUARY 2014 which vary with regards REFERENCES to the dominance of mosses, lichens, algae, Belnap, J., and O.L. Lange. 2003. Bio- logical Soil Crusts: Structure, Func- and fungi. In the Mojave tion, and Management, 2nd ed. Desert, significant differ- Springer Verlag, Berlin. ences in soil aggregate sta- Evans, R.D., and J.R. Johansen. 1999. bility, nitrogen fixation, Microbiotic crusts and ecosystem and carbon fixation can processes. Critical Reviews in Plant be detected among crust Sciences 18: 183–225. types. For example, li- Flechtner, V.R., N. Pietrasiak, and L.A. chen and moss crust com- Lewis. 2013. Newly revealed diver- munities contribute most sity of eukaryotic algae from wilder- to these ecosystem func- ness areas of Joshua Tree National tions, whereas some Park (JTNP). Monographs of the weakly-developed algal Western North American Naturalist 6:43–63. crusts perform poorly. Garcia–Pichel, F., et al. 2003. Small- Hence, the effect crusts scale vertical distribution of bacte- have on an area depends rial biomass and diversity in biologi- both on the community cal soil crusts from arid lands in the types present and how Colorado Plateau. Microbial Ecology– much of the ground is 46: 312–321. covered. Herrick, J.E., et al. 2010. Fine gravel In addition to the im- controls hydrologic and erodibility portant ecological role of responses to trampling disturbance these communities, they for coarse-textured soils with weak harbor a multitude of un- cyanobacterial crusts. Catena 83: discovered species and 119–126. genera. A research team Lovich, J.E., and D. Bainbridge. 1999. Anthropogenic degradation of the led by Valerie Fletchner SouthernCalifornia desert ecosys- investigated over 100 tem and prospects for natural recov- strains of eukaryotic al- ery and restoration. Environmental gae and discovered 15 Management 24: 309–326. clades (a clade is a group Pietrasiak, N., J.R. Johansen, and R.E. of organisms whose mem- Drenovsky. 2011. Geologic compo- bers share features from a sition influences distribution of mi- common ancestor), of crobiotic crusts in the Mojave and which only 6 could be Colorado Deserts at the regional identified to established scale. Soil Biology and Biochemistry genera. Recently, Radka 43: 967–974. considered to be the fertile mantle of Muehlsteinova, together with the Pietrasiak, N., et al. 2011. Spatial dis- tribution and comparison of distur- desert landscapes. authors, investigated over 150 cy- bance impacts to microbiotic soil The physiological performance anobacterial strains and found at crust in the Little San Bernardino of crust communities and the re- least 23 clades, with most of these Mountains of Joshua Tree National sulting ecological impact in the land- lineages being potential new genera Park, California. Western North scape varies with community com- and species to science. These find- American Naturalist 71: 539–552. position. Depending on climatic, ings are just an initial indication of Pietrasiak, N., et al. 2013. Biological geomorphic, and soil properties, how little we know about the soil crust community types differ in crust communities differ in the struc- biodiversity of these amazing mi- key ecological functions. Soil Biol- tural components of microbiotic croscopic communities. ogy and Biochemistry 65: 168–171. crusts (lichens, mosses, fungi, In summary, one of the greatest cyanobacteria). Where this happens, anthropogenic threats to desert sys- Nicole Pietrasiak and Jeffrey R. Johansen, crust communities have distinct tems is the loss of biological soil Biology Department, John Carroll Uni- forms and structures. The Great Ba- crust communities, and the con- versity, 1 John Carroll Boulevard, Univer- sin, Mojave, and Sonoran Deserts comitant loss of biodiversity, ero- sity Heights, OH 44118, npietrasiak@jcu. are especially rich in crust types, sion control, and soil fertility. edu, [email protected]

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 19 Although smaller streams may not always support classic desert wash plants, the denser and more robust vegetation along even smaller streams provides shade, escape cover, breeding and nesting sites, seed, and other food sources. Unnamed distributary channel on desert pavement near El Centro, CA. Photograph by Kris Vyverberg, California Department of Fish and Wildlife. DESERT STREAMS: AN INNOVATIVE NEW APPROACH TO THEIR INVENTORY AND MAPPING by Carolyn Chainey-Davis

One of the most startling paradoxes streams—driven by wildly unpre- method for delineating episodic of the world’s drylands is that al- dictable rain patterns, heavy flood streams in the California desert suc- though they are lands of little rain, flows, and sparsely vegetated and cessfully meets that challenge where the details of their surfaces are erodible soils—have a form and hy- other methods have failed. mostly the products of the action drology that are remarkably differ- Surprisingly, one of the most of rivers. To understand the natu- ent from streams elsewhere in the prominent features of desert land- ral environments of drylands is to state. scapes is their system of stream chan- understand the process and forms Concepts and methods used to nels (Schwinning et al. 2011). In- of their rivers. delineate streams that were devel- deed, sometimes entire alluvial —W.L. Graf (1988) oped for streams in areas of greater fans—those vast triangular aprons rainfall, and an historical bias for of sand, gravel, and arid soils that perennial streams, has led to an un- are built from arid mountain can- A PARADOX AND A derestimation of the extent and im- yons and comprise 30–40% of the PARADIGM SHIFT portance of desert streams. With the desert landscape—are characterized increasing pressure to develop re- by complex networks of actively he physical processes that newable energy projects in the shifting stream channels. These wa- shape streams in the desert desert, there is an urgent need for ter-driven landscape features breathe are very different from those an entirely new approach to the way life into otherwise harsh environ- in other parts of California. we perceive, map, inventory, and ments, but are routinely ignored or TheT hydrology of desert streams evaluate desert streams, one that rec- undervalued in development plan- has been likened to an early descrip- ognizes and incorporates their ning and environmental impact as- tion of life in the trenches of World unique physical, hydrological, and sessments. War I: “long periods of boredom, ecological processes. A new science- Until recently, change came brief moments of terror.” Desert based geomorphic and ecological slowly to the California desert. Habi-

20 FREMONTIA VOL. 42, NO.1, JANUARY 2014 tats were generally intact or mini- and behave differently than mally impacted, with disturbance perennial streams, episodic generally limited to highways, rail- streams provide all of the road and utility corridors, scattered same ecosystem services— mining operations, and sheep graz- ecological, hydrological, ing. Populations of many of the and physical (Levick et al. desert’s rare plants and animals were 2008). However, our cul- considered relatively stable. But in tural bias for perennial the last decade the political and eco- streams has led to a dis- nomic push for renewable energy missal of the desert’s epi- development has placed many desert sodic streams and the species and ecosystems at risk, eco- unique habitats they sup- systems inextricably linked to the port as having little or no functional integrity of streams and habitat value. their floodplains. The fundamental im- portance of ephemeral and Distributary channel networks on alluvial fans running off the Avawatz Mountains near Death Valley, CA. ECOSYSTEM BENEFITS intermittent streams to wildlife in the desert is un- They provide groundwater recharge and storm runoff to playa lakes and replenish the sand source critical All streams are life giving, but disputed. Streams and their for the maintenance of dunes. Oblique view of Google perhaps nowhere more so than in floodplains provide not Earth imagery provided by Jeremy Lancaster, California deserts. Streams occur in a variety only critical wildlife habi- Geological Survey. of channel forms in the Mojave and tat, but provide a founda- Sonoran Deserts, but there are very tion for much of the desert’s biotic of the Mojave River; or the distinc- few perennial streams (those that diversity. Higher soil moisture con- tive shrubby thickets of catclaw flow year-round). The most com- tent and topographic relief of desert (Senegalia greggii) or desert almond mon are the episodic streams, which streams provide shade and cooler (Prunus fasciculata) on the single- include ephemeral streams (flowing temperatures, which are used ex- thread channels of desert mountain briefly during and after a rain event) tensively by wildlife, and especially canyons. and intermittent streams (those that by less mobile creatures that cannot In comparison to larger streams, flow seasonally). Although they look avoid the harsh environment out- the smaller channels of alluvial fans side the stream ecosystem. The and their broad active floodplains higher moisture content, gravel, may not be as floristically distinct sand, and soils delivered by stream from adjacent plant communities. flows, and the rapid drainage prop- Yet they are nearly always measur- erties of these loose, stream-depos- ably denser, more robust, and have ited sediments support plant com- greater vegetative cover than the munities that are distinct from those adjacent drier uplands. The smaller on upland soils. This contrast is par- channels play an equally important ticularly stark on desert pavements role in seed dispersal and germina- where in some areas nearly all the tion, including the seeds of some wildlife habitat is found along rare species. The hard seed coats of streams, even in the smallest of some species, especially the legumi- channels. nous plants like ironwood, palo The streamside vegetation of verde, catclaw, and smoketree, are larger ephemeral streams can be abraded by the flows, promoting quite distinct: the floriferous wood- more efficient germination in seeds lands of palo verde (Parkinsonia that may otherwise require years of floridum), ironwood (Olneya teso- weathering to germinate. ta), and smoketree (Psorothamnus Where small streams dissipate Pooled water at the terminus of a small spinosus) that characterize the large on the valley floor, water sometimes channel where it dissipates near the toe of compound channels of the Sonoran pools at the terminus of the chan- an alluvial fan, three days after a quarter- Desert region of California; or the nel, and provides valuable tempo- inch summer storm event. Two weeks later, the channel was filled with native summer oases of lush Fremont cottonwood rary access to water for wildlife. annuals. Pahrump Valley, CA. Photograph (Populus fremontii) and sandbar wil- Pooled water and fine sediments can by Carolyn Chainey-Davis. low (Salix exigua) on wide reaches germinate abundant annuals follow-

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 21 Unnamed stream east of Twentynine Palms, San Bernardino County, California. The smaller swales that dissect the upper fan surfaces collect runoff and initiate stream flow from the surrounding, less permeable dark desert pavement surfaces. These are important sources of water, sediment, nutrients, and other materials during runoff, and are considered integral parts of a desert stream system. Photograph by Jeremy Lancaster, California Geological Survey. ing summer storms, like Bigelow’s food sources for desert tortoise and invertebrate resources and resting monkeyflower (Mimulus bigelovii), other species because they support opportunities for migrating shore- scarlet lupine (Lupinus concinnus), more diverse plant communities birds on their journey across the and shaggyfruit pepperweed (Lepi- with greater cover and higher diver- desert (Robinson and Oring 1996). dium lasiocarpum). These provide sity of summer and fall annuals. They fresh young greens for habitat and also host greater bird abundance and forage in a dry summer landscape. diversity, in fact up to 1.5 times NEW METHODS TO MAP Reptiles and amphibians are more breeding species and twice as DESERT STREAMS present in higher diversity in desert many wintering species than the streams than uplands, and many drier adjacent desert scrub (Kubick As pressure to develop the desert species preferentially use streams and Remsen 1977; Tomoff 1974; increases, it has quickly become and their floodplains because of Daniels and Boyd 1979a, 1979b). apparent that 1) the old concepts denser plant cover (Baxter 1988). The ecosystem benefits provid- and methods used to delineate Desert streams provide important ed by desert streams extend well streams in wetter regions—which beyond watercourse boundaries. assume regular and reliable flows, Downstream and downwind dune well-defined channels, or the pres- ecosystems are wholly dependent on ence of classic riparian zones— the replenishment of sands deliv- don’t work in the desert; and 2) ered by episodic streams to the there is an urgent need to develop “Aeolian” (wind-based) sand trans- a reliable and consistent method port systems that maintain dunes for mapping and evaluating streams and the many rare animals and plants that incorporates an understanding they support, such as Coachella of the distinctive physical, hydro- Valley milk-vetch (Astragalus len- logic, and ecological processes of tiginosus var. coachellae). Episodic desert streams. streams provide storm water runoff In response, the California De- to playa lakes and recharge the partment of Fish and Wildlife groundwater sources, which in turn (CDFW), with additional funding support the unique and often rare from the California Energy Com- Dense stringers of galleta grass (Hilaria habitats that occur at the playa mar- mission (CEC), is preparing a sci- rigida) line the small channels of an alluvial gins: marshes, saltgrass meadows, ence-based methodology for the de- fan on the Palo Verde Mesa near Blythe, alkali sink scrubs, and dune thick- lineation of desert streams to assist California, providing predator protection and a cooler microclimate for wildlife. ets. Where the playas and wetlands agency staff and energy project de- Photograph by Roland Brady, Brady and occur along migratory pathways, velopers, and to help interested par- Associates Geological Services. they seasonally provide critical ties in their review of project im-

22 FREMONTIA VOL. 42, NO.1, JANUARY 2014 pacts to streams. Though still in its dewatered portions of the fan, and No. 150. Paloverde desert wash. infancy, MESA: Mapping Episodic shrub density was reduced over 30%. American Birds 33(1): 94. Stream Activity (Vyverberg and Brady Such studies provide an important Graf, W.L. 1988. Fluvial Processes in 2013) provides a photographic atlas new perspective on the function, Dryland Rivers. Springer-Verlag, Ber- of the geomorphic indicators of epi- complex structure, and intrinsic lin, Germany. Johnson, H.B, F.C. Vasek, and T. Yon- sodic stream activity, mapping guid- value of distributary channel net- kers. 1975. Productivity, diversity, ance, guidance in the interpretation works, regardless of their size or the and stability relationships in Mojave of aerial photos, and six stream map- frequency of their flows. The desert Desert roadside vegetation. Bulletin ping case studies. Even to the non- is an enchanting place, home to many of the Torrey Botanical Club 102: 116– professional this field guide, which remarkable plants and animals that 115. is filled with interesting photos and are wonders of adaptation. The en- Kubik, M.R., Jr., and J.V. Remsen, Jr. diagrams, provides a riveting crash during health of these species and 1977. Fortieth breeding bird census. course in desert fluvial geomorph- their living habitats are undeniably No. 125. Catclaw-rabbitbrush desert ology (the shaping of rivers and linked to the functional integrity of wash. American Birds 31(1): 75–76. streams). This document will be the desert’s complex drainage sys- Levick, L., et al. 2008. The Ecological available on the CEC and CDFW tems. The challenge for the conser- and Hydrological Significance of Ephemeral and Intermittent Streams websites by spring 2014. vation, management, and restoration in the Arid and Semi-Arid American of the desert’s unique episodic stream Southwest. U.S. Environmental Pro- ecosystems begins with our percep- tection Agency and USDA/ARS STREAM INFLUENCE FAR tion of them. They are highly dy- Southwest Watershed Research BEYOND THE CHANNEL namic systems in time and space and Center. EPA/600/R-08/134, ARS/ their influence, particularly on allu- 233046. Washington, DC. Stream channels are among the vial fans, spans the entire watershed. Ludwig, J.A. 1986. Primary produc- most conspicuous features on the Regardless of the duration of tion variability i5n desert ecosys- desert landscape and contribute to their flows, episodic streams pro- tems. In Pattern and Process in Desert the ecological health of desert plant vide all the same ecosystem benefits Ecosystems, ed. W.G. Whitford. Uni- versity of New Mexico Press, Albu- communities (Schwinning et al. of perennial streams, and their eco- querque, NM. 2011). Thus, when we modify or logical importance is particularly Robinson, J.A., and L.W. Oring. 1996. eliminate desert streams we essen- significant given their vast extent. Long-distance movement by Ameri- tially halt life-giving flow to an MESA: Mapping Episodic Stream Ac- can avocets and black-necked stilts. entire watershed. For example, tivity by Vyverberg and Brady (2013) Journal of Field Ornithology 67: 307– multiple studies that compare the is an important starting point in the 320. vegetation of unaltered alluvial fans development of a new science-based Schlesinger, W.H., and C.S. Jones. to the dewatered portions down- method for mapping, inventorying, 1984. The comparative importance stream of a diversion have found and evaluating desert streams that of overland runoff and mean annual compelling results. No matter how incorporates an understanding of rainfall to shrub communities of the Mojave Desert. Botanical Gazette small the channels nor the frequency their unique forms and processes. 145: 116–124. or duration of flows, the health and Schwinning, S., et al. 2011. The influ- integrity of alluvial fan ecosys- REFERENCES ence of stream5 channels on distri- tems depends on the integrity of butions of Larrea tridentata and Am- their stream channel networks Baxter, R.J. 2008. Spatial distribution brosia dumosa in the Mojave Desert, (Schwinning et al. 2011). of desert tortoises (Gopherus agassi- CA, USA: Patterns, mechanisms, and Ecological impacts and the im- zii) at Twentynine Palms, California: effects of stream redistribution. pacts on stream-forming processes Implications for relocations. In Man- Ecohydrology 4(1): 12–25. are suffered far beyond the original agement of Amphibians, Reptiles, and Tomoff, C.S. 1974. Avian species diver- disturbance sites (Schwinning et al. Small Mammals in North America. sity in desert scrub. Ecology 55: 396– 403. 2011; Schlesinger and Jones 1984; Proceedings of the Symposium, July Vyverberg, K.A., and R. Brady. 2013. Ludwig 1986; Johnson et al. 1975). 19–21, 1988. US Forest Service Gen- eral Technical Report RM-166. Flag- MESA: Mapping Episodic Stream In one study near Joshua Tree Na- staff, AZ. Activity. California Department of tional Park (Schlesinger and Jones Daniels, B.E., and J. Boyd. 1979a. Fish and Wildlife. September 2013 1984) that compared vegetation on Forty-second breeding bird census. draft (unpublished). Sacramento, transects of 4,350–4,750 feet above No. 142. Ironwood-smoketree desert CA. and below a railroad that bisects an wash. American Birds 33(1): 92–93. alluvial fan, creosote bush suffered a Daniels, B.E., and J. Boyd. 1979b. Carolyn Chainey-Davis, 182 Grove Street, 50% reduction in biomass on the Forty-second breeding bird census. Nevada City, CA 95959 [email protected]

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 23 Desert riparian areas are relatively rare, yet provide critical breeding and migratory habitat for many endangered bird species. Photograph by Andrea Jones.

INSETS: (TOP) An endangered desert bird, the Least Bell’s Vireo, depends on desert washes and riparian habitats to breed. Photograph by Steve Mlodinow. • (MIDDLE) The Verdin, distinguished by its yellow facial pattern, is often found in desert areas containing desert washes. Photograph by Greg Smith. • (BOTTOM) Le Conte’s Thrasher is a bird that breeds in desert scrub and desert wash habitat in Southern California, including the Carrizo Plain. Photograph by Alan Schmierer. THE ECOLOGICAL IMPORTANCE OF WASHES TO DESERT BIRDS by Andrea Jones and Garry George

esert riparian habitats in (2000), Mark Dimmitt says that “dry includes significant stands of mi- Southern California are washes occupy less than 5% of this crophyll woodland associated with rare and of additional con- subsection of the Sonoran Desert, desert washes that flow east into servation concern because but support 90% of its bird life.” The the Colorado River, including the Dthey represent critical habitat for en- structural diversity in desert washes Chemehuevi Wash, Vidal Wash, and dangered breeding bird species such can include trees such as palo verde Miltiptas Wash. Another major oc- as Southwestern Willow Flycatch- (Parkinsonia aculeata) and ironwood currence of microphyll woodland ers, Western Yellow-Billed Cuckoos, (Olneya tesota) that provide breed- habitat lies at the base of the and Least Bell’s Vireos. In addition, ing and feeding habitat for a variety Algodones Dunes, halfway between in vast desert regions, these green of songbirds, particularly Phaino- the Colorado River and the south- patches with fresh water support peplas, Ash-Throated Flycatchers, ern end of the Salton Sea. In spring, thousands of songbird migrants Verdin, Crissal, LeConte’s and Ben- these areas attract large concentra- while they are en route from their dire’s Thrashers, Long-Eared and tions of migratory songbirds and wintering grounds in Central and Western Screech Owls, Black-Tailed provide nesting areas for breeding South America to breeding grounds Gnatcatchers, Gila and Ladder- species such as the Elf Owl, Gila further north. Backed Woodpeckers, Lucy’s War- Woodpecker, Long-Eared Owls, and Of particular note to avian com- blers, Northern Mockingbirds, and Bendire’s Thrashers. Additional in- munities are desert washes—dry Loggerhead Shrikes, some of which formation is available at: http://www. streambeds subject to rapid flow are California bird species of special prbo.org/calpif/htmldocs/desert. during flash flooding—in the Colo- concern. htm. rado and Mojave Deserts, which oc- The National Audubon Society cur in both upland and riparian habi- has recognized the significance of Andrea Jones, Suite 14, 601 Embarcadero, tat, and may include trees, shrubs, these habitats by naming the “Colo- Morro Bay, CA 93442, ajones@audubon. or a combination of both. In A Natu- rado Desert Microphyll Woodland” org; Garry George, 926 S. Citrus Ave, Los ral History of the Sonoran Desert an Important Bird Area. This area Angeles, CA 90036, [email protected]

24 FREMONTIA VOL. 42, NO.1, JANUARY 2014 JOHN O. SAWYER, JR.: 1939–2012 by James P. Smith, Jr.

ohn Sawyer passed away at his celled. It was there that students or Todd Keeler- CHRISTY NAVARRO home in Arcata on August 19, participants in CNPS or Jepson Her- Wolf, and Julie Evens, of the second 2012. His loss is deeply felt by barium field trips experienced his edition of A Manual of California Veg- Jhis wife, Jane Cole, his family, passion and detailed knowledge of etation. It is recognized as the stan- and his many colleagues and friends plants. He was not known for lead- dard reference on the subject. with whom he had close professional ing casual strolls across the coun- John wrote sections for Terres- and personal relationships. We will tryside. The lack of a well-defined trial Vegetation of California on the miss his comradeship, infectious trail was only a slight impediment if montane and subalpine vegetation laugh and smile, support, boundless there was a manzanita or a goose- of the Klamath Mountains, the for- energy, and wealth of knowledge. berry to be examined. ests of northwestern California, and With his passing, our state has lost John’s mobility became increas- alpine vegetation. His Trees and one of its most respected and influ- ingly limited by the effects of the Shrubs of California, coauthored with ential plant ecologists. The Califor- cancer that would take his life. So John Stuart, a Humboldt State col- nia Native Plant Society has lost a on our private sojourns together, league, has become a favorite field former president, an effective advo- we just changed our mode of opera- guide. He also produced a book with cate for the preservation of our na- tion to day trips in the car. John Andrea Pickart, a former graduate tive flora, and an individual who con- would spot a plant, usually of the student, on the ecology and restora- tinued to make significant contribu- woody persuasion, we would screech tion of coastal dunes in Northern tions to the society for over 40 years. to a stop, and Jane and I would California. John earned his bachelor’s degree gather specimens for him to look at. Some plant ecologists have little at California State University, Chico, His countless hours in the field, knowledge of the flora itself. Not so and his MS and PhD in plant ecology extending well beyond his formal with John. He was involved with from Purdue University. While pur- retirement, resulted in more than Dale Thornburgh in documenting a suing his graduate degrees, he was 40 books and papers! There were range extension of subalpine fir into an instructor at CSU, Chico, a gradu- excursions into such diverse topics California; authored a paper on the ate assistant at Purdue, and a field as the biological formations of the serpentine flora of the Lassics; co- ecologist for a consulting firm with eastern United States, the vegeta- authored the description of a new projects in Costa Rica and Thailand. tion life zones of Costa Rica, and subspecies of Frangula; and authored Sawyer joined the faculty of even a flora of a region in northern or coauthored four family treatments Humboldt State University in 1966 Thailand. But John’s first where he remained until his retire- passion was the vegeta- ment in 2001. He taught general ecol- tion of California, par- ogy, plant ecology, arctic and alpine ticularly that of the ecology, California natural history, northwestern part of the and plant taxonomy lab sections. state. Having gone into Early on, he was the prime mover in the field with him for 43 a very popular spring break offer- years, I can attest to both ing—the desert field trip—an inten- his boundless enthusi- sive excursion in a state bus from asm and incredible Arcata to Southern California. In all knowledge. of these classes John demonstrated In particular, as his critical thinking, and he insisted on students would say, John it in his students as well. He was had this thing for woody especially proud of his cadre of about plants, especially coni- 50 graduate students, many of whom fers. You can see this so remained close friends and who now clearly in his book, occupy professorships and respon- Northwest California: A sible positions around the country. Natural History. He was John out on the Bridge of Heaven trail in Colorado’s San John’s favorite instructional also a coauthor, along Juan Mountains, July 2001. Photograph by Todd Keeler- arena was the field, where he ex- with his close colleague Wolf.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 25 California. John was a frequent con- 2011 he received the J.C. Pritzlaff tributor to the CNPS Bulletin and Conservation Award from the Santa Fremontia. His first article, appear- Barbara Botanical Garden. He was ing in the CNPS Newsletter in 1973 active in several organizations, in- (later to become Fremontia), was on cluding as a Councillor of the Save the Klamath Region. A bibliography the Redwoods League and in the of John’s publications will soon be Ecological Society of America, where available at the Humboldt State Uni- he served as a technical advisor. versity Herbarium website (www. In addition to this impressive humboldt.edu/herbarium/). list of accomplishments, John was John was a founding member of also an avid hiker, serious photog- the North Coast Chapter of CNPS in rapher, world traveler, follower of 1970 and served as its first president. Purdue University football, habitué LEFT TO RIGHT: John Sawyer, Virginia Keeler- He was especially interested in the of art museums, student of the op- Wolf, and his wife Jane Cole, enjoying a ride in an outboard up the Yacumama River vegetation and flora of the Klamath era, and an admirer of the music of of the Amazon Basin, Peru, summer 2002. region of northwestern California and Bach, Villa Lobos, Piazolla, Ives, Photograph by Todd Keeler-Wolf. southwestern Oregon. His field trips Loreena McKennitt, Cecilia Bartoli, to the Lanphere Dunes, Russian Peak, Bob Dylan, and Judy Collins. in The Jepson Manual and the treat- The Trinity Alps, Scott Mountain, John would not have wanted a ment of Rhamnaceae that will ap- Stony Creek Bog, Walker Ridge, and traditional memorial service. In its pear in the Flora of North America serpentine areas were legendary. place, Jane, her sister Susan, Michael North of Mexico. His local conservation efforts Kauffmann (a fellow “conehead”), John and I wrote 5 editions of were exceptional. John was a criti- and I organized a celebration of our Keys to the Families and Genera cal player in the establishment of John’s life and accomplishments at and Vascular Plants in Northwestern the Lanphere Dunes near Arcata and Patrick’s Point State Park, just north California, and 22 editions of our in maintaining its natural state by of Arcata. About a hundred people checklist of the plants of that same organizing “lupine bashes” to eradi- attended. As you would expect, there region have appeared. We were work- cate the invasive yellow bush lupine was a time for recollections. I ing on a specimen-based checklist of (Lupinus arboreus). The coastal thought that I knew John pretty well, the vascular plants of the Klamath dunes were also the home of one his but I was amazed at stories that were Region, and Sawyer exacted a prom- major research efforts on the status new to me, at the depth of emotion ise that I would see it completed. of the Menzies’ wallflower (Erysi- expressed, and at how important he John became the fourth presi- mum menziesii). had been in the lives of so many dent of CNPS in 1973, the first “out- He was also intimately involved who were there and others who sent sider” to occupy the position, in the in the work that led to the designa- remembrances to be read. sense of not being one of the group tion of the Russian Wilderness in If I might end on another per- that founded the society. He began Siskiyou County. There he and Dale sonal note, someone once observed the process of making CNPS a state- Thornburgh would document the that you are lucky in life if you have wide organization. Board meetings occurrence of 17 conifers in a one- one really good friend. If so, I was that had been held on Thursday eve- square-mile area around Little Duck very lucky indeed. nings, typically in the home of a Bay Lake, which must make it one of the Memorial contributions may be Area member, were moved to Satur- richest conifer forests in the world. made to any of the organizations day. An unfinished set of bylaws John and Jane assisted our friend mentioned above, to a memorial that had not permitted chapter presi- and colleague, Dave Imper, in pro- bench that will be dedicated to John dents to vote at board meetings was tecting serpentine barrens in The at the Lanphere Dunes (Friends of rewritten. He would later serve as a Lassics (Humboldt and Trinity coun- the Dunes, P.O. Box 186, Arcata member of five state-level commit- ties), the home of the endemic 95518), or to the John O. Sawyer, Jr. tees: Conservation, Educational Lassics lupine (Lupinus constancei). Endowment to support field studies Grants, Publications, Rare Plants, Among his many honors, John in botany (Advancement Founda- and Vegetation. I doubt that anyone was recognized by Humboldt State tion, Gift Processing Center, else can match that record of such University as its 1997 Scholar of the Humboldt State University, 1 Harpst diverse committee service. Year, by the California Botanical So- Street, Arcata 95521-8299). He was an editor of the second ciety in the 2008 dedication of its edition of the CNPS Inventory of Rare journal, Madroño, and by CNPS in James P. Smith, Jr., 884 Diamond Drive, and Endangered Vascular Plants of making him a Fellow in 1994. In Arcata, CA 95521, [email protected]

26 FREMONTIA VOL. 42, NO.1, JANUARY 2014 BOOK REVIEWS

I Am the World’s Oldest-Known Living Thibault takes the opportunity to teach Living Tree, teach- Tree by Dr. Gilbert G. Thibault. 2012. good science rather than avoid topics ers and parents can Winmark Communications, Glendale, contentious to a few. There are several use the text to meet AZ. 56 pp, $21.56 hardcover. ISBN# references to the potential impact of the goals of the 978-1-89222-515-3. climate change on the bristlecones. The state’s curriculum author notes that they are threatened in standards in science This book is dedicated “to the chil- summer by bark beetles, which die off literacy, particu- dren of the world and their parents . . . in cold weather. A rise in global tem- larly in grades 3–6. and to the wise people who had the peratures may endanger them and ben- They expect students will “actively foresight to preserve and protect this efit the bark beetle, since warmer tem- seek the wide, deep, and thoughtful unique habitat. . . . ” This fine 2012 peratures encourage the development engagement with high-quality literary children’s book about bristlecone pines of softer wood, which bark beetles pre- and informational texts that build will delight all readers and deepen their fer. On the other hand, there is a missed knowledge, enlarge experience, and appreciation of this remote native. opportunity to explore balanced eco- broaden worldviews.” This book cer- From the first leaf, author/photog- logical relationships: pine bark beetles tainly qualifies as such a text. rapher Gil Thibault’s graphic expertise are eaten by Clark’s Nutcrackers, which Author Gil Thibault has written a is evident. Soft greyed illustrations point disperse bristlecone seeds. In nearby volume that’s full of wonder, visual out detail the reader might miss in the yellow pine forests, bark beetles are a and verbal treasures, accurate science, striking photos, giving the reader the normal part of fire ecology. humor, and reverence for bristlecones ability to see the forest and the trees. A In an adaptation that improves on and their habitat. I Am the Oldest Known tone of respect for these trees is set in Joyce Kilmer’s sing-song cadence, Living Tree makes one want to ensure the beautifully detailed introductory Thibault adapts his well-known poem that these venerable trees survive for map. Section themes pique curiosity, “Trees” to celebrate the bristlecone. generations to come. particularly the bristlecone as a weather Also included is a folksong, “Bristle- —Barbara Roemer and indicator, historian, or survivor. Stun- cone Pine,” by Hugh Prestwood, along Merry Byles-Daly ning photos of bristlecone pines against with an online video link. The lyrics the night sky are accompanied by an evoke the forest, but mentions of Jesus invitation to visit them, and an assur- and heaven and hell may put off non- CORRECTION ance that “ . . . we will leave the lights of Christian readers. Along the same trail, the Milky Way on for you.” the timeline would benefit from more The photo caption on p. 2 of Fremontia, Vol. 41, No. 3 (September 2013) iden- A master storyteller, Thibault en- non-Western entries. Small criticisms tified Fremont Canyon as located in gages the reader in devising names for in a stunning book. Irvine County. It is actually located near creatures and trees and speculating While a broad audience will be the city of Irvine, but in Orange County. about phenomena. He informs with fas- captivated by I Am the Oldest Known cinating facts: scientists have found downed deadwood that’s 11,000 years old, and the trees do not die of old age! Humor threads through nearly every page: “I have heard some people say my foliage is very green for my age.” Chil- dren will love the extremes he describes, from stormy winter weather to drought. Told from the perspective of The Methuselah, a 5,000-year-old bristle- cone pine, the book is unabashedly anthropomorphic. And in an interest- ing variant of personification, Methu- selah suggests humans can learn adapt- ability and resilience from bristlecone pines. A fascinating four-page timeline places these trees amid human habita- tion, discovery, and invention. Children of all ages will be drawn to the rich imagery the scientific lan- guage elicits, from dendrochronology, contiguous, and fortitude, to dolomite.

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 27 The plant guide that follows is orga- nine insect species associated with Wildflowers of Orange County and the nized alphabetically by family and, milkweed and dogbane are described Santa Ana Mountains by Robert L. Allen within family, more loosely by genus (with photographs!) for the reader. The and Fred M. Roberts, Jr. 2013. Laguna and taxonomic relatedness. A tremen- etymology of all genus and species Wilderness Press, Laguna Beach, CA. dous amount of information has been names is provided. Symbols are added 498 pages, $35.00, soft-cover. ISBN packed into the book; literally no inch for sensitive species that have federal 978-0-9840007-1-5. of space is wasted and nearly every or state protected status, as well as for species is not only succinctly described, those of current local concern. Southern California but also represented by multiple pho- While the guide is extremely thor- plant enthusiasts—espe- tographs. ough, readers should note that it is not cially those in Orange Even local variations within a spe- all-inclusive. A scattering of less showy County—have reason to cies such as occur in California buck- species such as walnut, oak, alder, celebrate the publication wheat (Eriogonum fasciculatum), coast members of the goosefoot family, and of a new book. Wildflow- goldenbush (Isocoma menziesii), and all grasses and their relatives are ex- ers of Orange County and gum plant (Grindelia camporum) are cluded to save space. Several non-na- the Santa Ana Mountains, diagrammed or described with photo- tive plants occurring in wildland habi- by Robert (Bob) Allen graphs. Key anatomical and morpho- tats are also missing. Keeping both and Fred Roberts, is a logical traits such as floret structure in would have moved the book out of the glorious nearly 500-page sunflowers or flower keel shape and realm of a field guide and into that of a guide to the showy flora distribution of hairs in lupines are also desk reference. of Orange County and clearly diagrammed to allow for easier Most refreshing to me is that the adjacent areas. identification. authors take a holistic view of plants Nearly ten years have passed since Allen and Roberts’ book has sev- in our fragile landscapes. What they rumors of this ambitious project first eral other unique and refreshing fea- have produced will not only serve as a surfaced, and the end product will thrill tures that place it in the top tier of local field guide to specialists and lay people, readers. The authors are two child- natural history references. Pollinators but also as an encyclopedia of local hood classmates from Orange County and animal guilds are described for plants and their ecology. who evolved along parallel paths as key species and habitats. For example, —Jutta Burger botanists and natural historians. Bob Allen, known to his followers as “Bug Bob,” is well known for his extensive local knowledge of plants and insects, as well as his entertaining public pre- Telos Rare Bulbs sentations and classes. Wildflowers be- The most complete offering of bulbs native to the gan as his brain child. Fred Roberts is western USA available anywhere, our stock is the leading authority on status and propagated at the nursery, with seed and plants distribution of plants in Orange County from legitimate sources only. and has enriched the county as botani- cal guru and artist for years. Together, they are the perfect team to have cre- ated this work. Telos Rare Bulbs The authors first introduce the re- P.O. Box 1067, Ferndale, CA 95536 gion, habitat types, and terminology www.telosrarebulbs.com needed to identify flowering plants.

28 FREMONTIA VOL. 42, NO.1, JANUARY 2014 Join Today! CNPS member gifts allow us to promote and protect California’s native plants and their habitats. Gifts are tax-deductible minus the $12 of the total gift which goes toward publication of Fremontia. ❏ $1,500 Mariposa Lily ❏ $600 Benefactor ❏ $300 Patron ❏ $100 Plant Lover ❏ $75 Family ❏ $75 International or Library ❏ $45 Individual ❏ $25 Limited Income CORPORATE/ORGANIZATIONAL ❏ $2,500 10+ Employees ❏ $1,000 7-10 Employees ❏ $500 4-6 Employees ❏ $150 1-3 Employees

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SUBMISSION INSTRUCTIONS CNPS members and others are invited to submit articles for pub- lication in Fremontia. If inter- ested, please first send a short summary or outline of what you’d like to cover in your ar- ticle to Fremontia editor, Bob Hass, at [email protected]. Instruc- tions for contributors can be found on the CNPS website, www.cnps.org, under Publica- tions/Fremontia.

Fremontia Editorial Advisory Board Susan D’Alcamo, Jim Andre, Ellen Dean, Phyllis M. Faber, Holly Forbes, Dan Gluesenkamp, Brett Hall, David Keil, Pam Muick, Bart O’Brien, Roger Raiche, Teresa Sholars, Dick Turner, Mike Vasey

VOL. 42, NO. 1, JANUARY 2014 FREMONTIA California Native Plant Society Nonprofit Org. 2707 K Street, Suite 1 Sacramento, CA 95816-5130 U.S. Postage PAID A.M.S.

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CONTRIBUTORS FROM THE EDITOR

James M. André is director of the Granite Mountains Desert his issue and the next of Fremontia, two issues focused Research Center, senior advisor to the CNPS Rare Plant on California’s deserts, exemplify what CNPS does Program (RPP), and chair of the RPP Committee. best. As the leading organization concerned with the healthT of the state’s native plants and vegetation types, we Duncan Bell is a field botanist for Rancho Santa Ana Botanic Garden, and spends most of his time in the undercollected bring together in one place science-based information and unexplored mountain ranges of Southern California. gleaned from the work of our own staff and a cadre of CNPS Jutta Burger, PhD, is the managing director of the science citizen-scientists, along with that of other prominent scien- and stewardship department at the Irvine Ranch Conservancy tists from academia, government agencies, and independent in Irvine, California. consultants. This information is then available to guide Merry Byles-Daly and Barbara Roemer are members of those who are entrusted to make decisions about how best the Redbud Chapter of CNPS and elementary educators. to manage and conserve our environment. Together they have logged over 60 years in the classroom. For those who know little about California’s deserts, Carolyn Chainey-Davis is a botanical consultant who these two issues will be eye-openers, and are likely to dispel recently conducted analyses of the biological impacts of six the myth that a desert area is a wasteland devoid of ecologi- proposed solar power plants in the California desert. cal value. One of the reasons some may hold that view is Garry George directs the Chapter Network and renewable because the richness of deserts is not readily apparent to the energy programs for Audubon California. untrained eye. Much is hidden by nature to protect it from the comparative harshness of the desert ecosystem. But after Jeffrey R. Johansen is professor of biology at John Carroll reading the articles in this issue, you may be amazed at all University. you discover should you decide to take a hike in any of Andrea Jones directs the Coastal Stewardship Program for California’s deserts. Audubon California. We owe a debt of gratitude to those researchers who Patrick McIntyre is an ecologist with the Berkeley Initia- are willing to endure the challenges presented by deserts tive in Global Change Biology at UC Berkeley. His research in order to inform us of what is happening there. Research- includes predicting rare desert plant distributions. ers are a unique, unheralded lot, and their work demands Kara A. Moore is an ecologist in the Center for Population considerable patience, precision, dedication, and a system- Biology at UC Davis and studies the dynamics that influ- atic approach. Each adds their own piece to the jigsaw of ence species distributions and the persistence of rare plants. scientific knowledge. While it is exciting to discover new Nicole Pietrasiak is a post-doctoral researcher in the biol- species and occurrences and to work in beautiful surround- ogy department at John Carroll University. ings, most botanists I know are more concerned with pro- James P. Smith, Jr. is professor of botany, emeritus, and tecting our native plants and ensuring that their vitality and curator of the herbarium at Humboldt State University in beauty endures. Arcata, California. —Bob Hass

FREMONTIA (continued on inside back cover) VOL. 42, NO. 1, JANUARY 2014