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VOL. 41, NO. 3 • SEPTEMBER 2013 FREMONTIA JOURNAL OF THE NATIVE SOCIETY

THETHE FUTUREFUTURE OFOF CALIFORNIACALIFORNIA SAGESAGE SCRUBSCRUB THETHE ARICAARICA MOUNTAINSMOUNTAINS LIFELIFE INSIDEINSIDE PLANTPLANT GALLSGALLS THE NEED FOR SITE-SPECIFIC MAINTENANCE MANUALS VOL. 41, NO. 3, SEPTEMBER 2013SEQUOIA:: CONTINUATIONCONTINUATION OF FREMONTIA THE SAGA 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. 41, NO. 3, SEPTEMBER 2013 MEMBERSHIP Copyright © 2013 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 or Group ...... $75 Benefactor ...... $600 International or Library ...... $75 Beth Hansen-Winter, Designer Patron ...... $300 Individual ...... $45 Brad Jenkins, Cynthia Powell, and Plant Lover ...... $100 Student/Retired/Limited Income . $25 Mary Ann Showers, Proofreaders CORPORATE/ORGANIZATIONAL + CALIFORNIA NATIVE 10 Employees ...... $2,500 4-6 Employees ...... $500 7-10 Employees ...... $1,000 1-3 Employees ...... $150 PLANT SOCIETY STAFF – SACRAMENTO CHAPTER COUNCIL Dedicated to the Preservation of Executive Director: Dan Gluesenkamp David Magney (Chair); Larry Levine the California Native Flora Finance and Administration (Vice Chair); Marty Foltyn (Secretary) Manager: Cari Porter Alta Peak (Tulare): Joan Stewart The California Native Plant Society Membership and Development Bristlecone (Inyo-Mono): (CNPS) is a statewide nonprofit organi- Coordinator: Stacey Flowerdew Steve McLaughlin Conservation Program Director: zation dedicated to increasing the Channel Islands: David Magney Greg Suba understanding and appreciation of Dorothy King Young (Mendocino/ California’s native , and to pre- Rare Plant Botanist: Aaron Sims Vegetation Program Director: Sonoma Coast): Nancy Morin serving them and their natural habitats Julie Evens East Bay: Bill Hunt for future generations. 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FREMONTIA VOL. 41, NO. 3, SEPTEMBER 2013 CONTENTS

THE FUTURE OF CALIFORNIA SAGE SCRUB IN AN ERA OF INCREASING URBANIZATION AND GLOBAL CLIMATE CHANGE by Erin C. Riordan and Philip W. Rundel ...... 2 How will the imperiled plant association, California sage scrub (CSS), fare under a future of increased habitat conversion from development and from climate change?

THE : BOTANICAL EXPLORATIONS OF A PHYTOGEOGRAPHICAL BLACK HOLE by Duncan S. Bell ...... 8 The Arica Mountains are an unexplored, rugged section of desert in Southern California that at first glance appear barren, but actually contain a wealth of biodiversity and numerous rare plants.

INDUCERS, PARASITOIDS, AND INQUILINES: LIFE INSIDE THE PLANT GALL by Peter Bryant ...... 14 When gall wasps or midges lay their eggs on an appropriate host plant, some of the plant cells are dramatically reprogrammed to produce amazing structures called galls that can be totally different from anything else that the plant ever makes.

THE NEED FOR SITE-SPECIFIC MAINTENANCE MANUALS FOR PUBLIC AND PRIVATE NATIVE PLANT LANDSCAPES by Ellen Mackey and Nancy L.C. Steele ...... 20 Site-specific maintenance manuals are more likely to ensure that native plant land- scapes thrive. Here’s how to create one.

SEQUOIA: CONTINUATION OF THE SAGA by Marcel Rejmánek ...... 26 G. Ledyard Stebbins and more contemporary researchers offer insights into the botanical origins of the redwood, one of the iconic trees of California.

THE COVER: Intact spiny bud galls induced by the wasp Diplolepis polita on a of California wild rose (Rosa californica). Photograph by Peter Bryant. Full story begins on page 14.

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 1 ABOVE: View of intact California sage scrub in spring bloom at Fremont Canyon in Irvine County, California. Photograph by Stephen Francis for the Irvine Ranch Conservancy. • FACING PAGE: View of California sage scrub in summer, dominated by California buckwheat (Eriogonum fasciculatum), a semi-evergreen species. All photographs by Erin C. Riordan unless otherwise noted. THE FUTURE OF CALIFORNIA SAGE SCRUB IN AN ERA OF INCREASING URBANIZATION AND GLOBAL CLIMATE CHANGE by Erin C. Riordan and Philip W. Rundel

alifornia sage scrub (CSS), patches along the central California sake aromatic sages (Salvia spp.), one of California’s most coast. In Southern California, it oc- and a rich understory of herbaceous threatened vegetation curs almost entirely in the wildland- native plants. types, faces a tenuous fu- urban interface in a complex and California sage scrub is rapidly Cture marked by rising threats from delicate mosaic of chaparral, grass- disappearing in California under a human land use and climate change. lands, and riparian corridors amidst variety of human pressures, and a This distinct vegetation type spans the region’s sprawling human popu- large proportion of species associ- coastal and semiarid interior regions lation. This vegetation lends a ated with CSS are categorized as sen- of the state from Santa Barbara south unique beauty to the Southern Cali- sitive, rare, threatened, or endan- to El Rosario in northwestern Baja fornia landscape with its drought gered. Land development, invasive California, as well as scattered deciduous shrubs, such as the name- species, changes in fire regime, and

2 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 air pollution have all contributed to ADDRESSING FUTURE environmental variables. Scientists the widespread loss, fragmentation, ENVIRONMENTAL CHANGE then use these models to map po- and degradation of CSS, which cur- tential habitat for a species under rently occupies a tiny fraction of its Incorporating scenarios of future future conditions. Species distribu- original extent (Westman 1981). environmental change into conser- tion models have been used for a Moreover, human activities that in- vation and resource management is variety of purposes, including to crease disturbance facilitate the con- challenging but increasingly neces- identify species at risk of habitat version of CSS to exotic grasslands, sary. Evidence of the negative eco- loss and extinction and to predict a change that is further reinforced logical ramifications of climate changes in biodiversity. by soil nitrogen inputs from air pol- change has accumulated over the In our study, we used a species lution. last decades, spurring a growing distribution modeling approach to To make the situation worse, CSS community of scientists to investi- predict changes in habitat for a num- faces additional threats from both gate the current impacts of climate ber of key CSS species under 21st projected global climate change and change and predict likely future im- century climate change. We used human population growth in South- pacts as the climate continues to location data from herbarium ern California. A rapidly warming warm. records to model the current cli- climate this century could cause spe- Unfortunately, California sage mate requirements for CSS species cies range shifts, species losses, and scrub exists inside a highly urban- (Riordan and Rundel 2009). Then declines in diversity as individual ized region with a growing human we mapped areas meeting species species respond to climate change, population. For this vegetation, fu- requirements under two possible tra- while increasing human populations ture land use may pose at least as jectories of projected climate change will likely cause additional loss of big a threat as climate change. Cali- in California: a warmer, wetter fu- wildlands. By the end of this cen- fornia is, of course, not the only ture and a warmer, drier future. tury, scientists expect California’s region facing such threats. It is one In order to weigh the impact of climate will shift substantially with of the world’s five Mediterranean future climate change against that of temperatures rising by 1.7–5.8 C climate regions, and in each one future human land use, we used pro- (3.0–10.4 F) statewide, depending human land use followed by climate upon the level of mitigation of green- change are expected to cause some house gas emissions (Mastrandrea of the highest biodiversity losses and Luers 2012). worldwide by 2100 (Sala et al. 2000). One study predicts dramatic Innovative modeling now makes range losses in as many as two-thirds it possible for scientists to predict of California’s endemic species by the potential impacts of future cli- 2100 (Loarie et. al 2008). Such find- mate change on biodiversity. The ings are particularly concerning for magnitude and trajectory of climate California, where endemic taxa change depends heavily upon hu- contribute greatly to the state’s bio- man greenhouse gas emissions, diversity, accounting for nearly 35% which will be determined by popu- of the 6,500 native taxa. California’s lation growth, economic develop- characteristic Mediterranean-type ment, technological development, climate of mild, wet winters and and land use. The Intergovernmen- warm, dry summers may be at risk tal Panel on Climate Change (IPCC) in coastal areas of Southern Califor- has released a set of widely used nia and northwestern Baja Califor- possible future emission scenarios nia (Ackerly et al. 2010; Klausmeyer based upon different patterns of hu- and Shaw 2009). Warming and dry- man growth and development. So- ing conditions could cause the loss phisticated global climate models are of Mediterranean-type conditions in then used to generate predictions of these areas, which harbor high CSS future climate patterns under these diversity and endemism. future emission scenarios. In the face of these mounting Species distribution models can changes, we set out to assess the provide hypotheses of how species potential impacts of both 21st cen- may respond to climate change. tury climate change and land use on These models define a species range CSS shrub species in California. with respect to climate and other

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 3 nomic and population growth in Southern California fueled extensive settlement and urbanization of low- land coastal areas, much at the ex- pense of the native shrublands such as CSS. Today, nearly half (45%) of California’s 38 million residents live in the coastal counties of Southern California (Ventura, Los Angeles, Orange, and San Diego), which to- gether comprise just 7% of the state’s total land area. High land values and development pressure across the re- gion pose a significant threat to re- maining fragments of CSS, making conservation both costly and chal- lenging. These pressures will con- tinue to grow with California’s popu- lation, which is projected to increase to as high as 147.7 million by 2100 (Sanstad et al. 2011). Approximately 25% of coastal and western Southern California has now been developed primarily as a result of urban and suburban growth, transportation infrastruc- ture, and agriculture. This percent- age could rise to over 40% by late TOP: Conversion of California sage scrub to century (2080s) under a future agricultural land uses in , marked by high economic growth Mexico. Photograph by Benjamin T. Wilder, and urbanization. Under such a sce- 2010. • BOTTOM: Black sage (Salvia mellifera) with a view of sprawling urbanization in nario, the single greatest type of land the Los Angeles basin outside of Pasadena, conversion in the region will be the California, 2010. loss of shrubland habitats to devel- opment. for the first time in Southern Cali- Such high rates of future habitat fornia. Our model of future CSS ex- conversion will likely impact future tent was based on a future of high CSS persistence. For the 20 domi- economic growth, high urbanization, nant species considered in our study, and high greenhouse gas emissions. we predict that species may lose, on average, 30% of remaining uncon- verted and suitable habitat in Cali- MOUNTING DEVELOPMENT fornia to land use alone by late cen- PRESSURES tury. We predict habitat losses of jections of future land cover and land greater than 40% in coastal species use released by the California sage scrub’s primarily with southern distributions, such as Geological Survey (USGS) Land Car- Southern California distribution co- the coast prickly pear (Opuntia bon project, the first-ever of such incides with areas of high human littoralis) and San Diego sunflower projections for the conterminous impact. (Bahiopsis laciniata). We predict ap- United States (Sleeter et al. 2012). Southern California’s history of proximately 20% of habitat conver- These projections, built upon the land use dates back to the Spanish sion for some of the more wide- same future emission assumptions ranchos of the late 18th century and spread species, such as California as the future climate projections, al- subsequent Euro-American settle- buckwheat (Eriogonum fasciculatum) low us to directly compare the two ment during the 19th century. Fol- and sticky monkeyflower (Mimulus main causes of environmental change lowing World War II, rapid eco- aurantiacus).

4 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 A FUTURE OF CLIMATE CHANGE By the end of the century, we predict that dominant CSS species will lose considerable habitat in Southern California due to climate change alone. If we assume that CSS species are unable to disperse and track favorable conditions outside of their current ranges, the species we studied may lose on average 30% of their remaining suitable habitat due to climate change alone, but the loss increases to 50–60% after we factor in habitat conversion under projected land use. Some species, such as mis- sion manzanita (Xylococcus bicolor), may lose up to 88% of suitable habi- tat due to land use and climate Characteristic CSS species, FROM TOP TO change. These predictions are par- BOTTOM: coast prickly pear (Opuntia littor- alis); sticky monkeyflower (Mimulus auran- ticularly troubling, given the high tiacus); California buckwheat (Eriogonum diversity and high number of threat- fasciculatum) and golden yarrow; and the ened species associated with CSS in San Diego sunflower (Bahiopsis laciniata). Southern California. Coast prickly pear and San Diego sunflower Along the Central California are predicted to experience large losses in habitat to human land uses by 2080. Sticky coast, however, we predict a strik- monkeyflower photograph by Michael W. ingly different pattern. Here, climate Tuma. San Diego sunflower photograph by change may cause the environment Benjamin Smith. to become more favorable for CSS shrubs. Assuming CSS species can nario), even after accounting for fu- spread to newly suitable areas, we ture land use change. These habitat predict species will experience an gains, much of which will occur in overall net gain of habitat in Califor- the Central Coast, could offset some nia by an average of 15% (in a of the habitat losses predicted in warmer, wetter future scenario) to Southern California, and cause a gen- 21% (in a warmer, drier future sce- eral shift northward in CSS distribu- tion. We predict consid- Under future climate scenarios, the evergreen shrub erable net habitat gains lemonade berry (Rhus integrifolia), is projected to gain in California of over climatically suitable habitat along California’s central coast 90% for three species: by late century (2080s), assuming species can disperse or spread into newly suitable areas. Photograph by Alan B. California bush sun- Harper. ( califor- nica), lemonade berry (Rhus integrifolia), and coast prickly pear (Opun- tia littoralis). Even assuming spe- cies can move and spread under climate change, California sage scrub will likely face sig- nificant obstacles to fu- tat. The considerable land develop- ture range expansion ment and habitat degradation that into areas of new habi- characterize coastal Southern Cali-

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 5 fornia pose formidable barriers to needed to mitigate habitat loss from The future persistence of CSS species movement. Competition both future land use and climate will also hinge upon the successful with already established species change. establishment of species along could also prevent CSS from colo- nizing areas of new habitat without FIGURE 1. FUTURE HABITAT GAIN UNDER CLIMATE CHANGE BY LATE additional human assistance. A mo- CENTURY (2080S) FOR CALIFORNIA BUSH SUNFLOWER (ENCELIA saic of chaparral, CSS, and grass- CALIFORNICA). lands dominate much of the central coast, with local patterns of CSS dis- tribution governed by a number of factors in addition to climate, such as fire, topography, soil type, and species interactions.

IMPLICATIONS FOR CONSERVATION As climate conditions change, the future persistence of species may well hinge upon their ability to move to areas with favorable environmen- tal conditions. Our models under- score the importance of dispersal in moderating CSS habitat losses. How- ever, human activities such as habi- tat conversion through land use and altered disturbance regimes pose a First panel shows current modeled suitable habitat (gray) and locations where the species major barrier to plants realizing their has been recorded. In future panels, blue represents habitat gain, red habitat loss, and gray full dispersal potential. Additionally, remaining climatically suitable habitat. Photograph by Erin C. Riordan. the factors influencing a species’ suc- cessful establishment are numerous FIGURE 2. SEVERE HABITAT LOSS UNDER CLIMATE CHANGE BY LATE and nuanced. Thus, the future dy- CENTURY (2080S) FOR MISSION MANZANITA (XYLOCOCCUS namics of CSS expansion will likely BICOLOR). be complex, governed by many pro- cesses that can also be influenced by human activities. When species respond individu- ally to climate change, we can ex- pect patterns of diversity to change as well. We predict considerable de- clines in CSS shrub diversity in coastal Southern California. Vegeta- tion composition will change, which could have implications for the large number of threatened and endan- gered animals that depend upon CSS for habitat. If CSS species are able to shift northward in response to cli- mate change, we can expect changes in vegetation along the Central Cali- fornia coast. In Southern California, remaining high quality CSS vege- tation needs to be protected from First panel shows current modeled suitable habitat (gray) and locations where the species development and new approaches has been recorded. In future panels, blue represents habitat gain, red habitat loss, and gray to resource management may be remaining climatically suitable habitat. Xylococcus bicolor photograph by Keir Morse.

6 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 FIGURE 3. COMPOUNDING IMPACTS OF FUTURE CLIMATE mate change may be more painful CHANGE AND LAND USE ON CALIFORNIA SAGE SCRUB SHRUB than the costs of curbing our carbon DIVERSITY. emissions. Hopefully grim predic- tions, such as those from our mod- eling effort, will spur action rather than grief or resignation.

REFERENCES Ackerly, D.D., et al. 2010. The geogra- phy of climate change: implications for conservation biogeography. Diver- sity and Distributions 16: 476–487. Klausmeyer, K.R., and M.R. Shaw. 2009. Climate change, habitat loss, protected areas, and the climate ad- aptation potential of species in Medi- terranean ecosystems worldwide. Plos One 4: e6392. doi:10.1371/ journal.pone.0006392. Loarie, S.R., et al. 2008. Climate change and the future of California’s en- demic flora. Plos One 3: e2502. doi:10.1371/journal.pone.0002502. Mastrandrea, M.D., and A.L. Luers. 2012. Climate change in California: Scenarios and approaches for adap- tation. Climatic Change 111: 5–16. Riordan, E.C., and P.W. Rundel. 2009. Modelling the distribution of a threatened habitat: The California sage scrub. Journal of Biogeography 36: 2176–2188. Modeled species richness ranges from 1 (blue) to 33 (red). Areas of habitat conversion to Sala, O.E., et. al. 2000. Biodiversity— human land uses in Southern California are shown in dark gray in the bottom three panels and overlap with areas of species loss driven by climate change. The coastal central Global biodiversity scenarios for western and southwestern California ecoregions are shown in a black outline. the year 2100. Science 287: 1770– 1774. coastal Central California. In this CSS may be impacted by future cli- Sanstad, A.H., et al. 2011. Projecting long-run socioeconomic and demo- case, the role for managers is more mate and land use changes. They graphic trends in California under complex. How actively should man- can best be applied in combination the SRES A2 and B1 scenarios. Cli- agers facilitate species movements? with careful monitoring of CSS and matic Change 109: 21–42. Does the northern expansion of CSS climate change and land use impacts Sleeter, B.M., et al. 2012. Scenarios of species come at the expense of local in an adaptive management context. land use and land cover change natives or other vegetation types, Conservationists—including in the conterminous United States: such as chaparral? One approach, CNPS members, land managers, Utilizing the special report on emis- which is not specific to CSS and is agency personnel, and environmen- sion scenarios at ecoregional scales. likely to benefit a variety of species, tal advocates—need to start think- Global Environmental Change 22: is to promote species movements ing about whether it is proper to 896–914. through the protection of strategic allow and/or facilitate the expansion Westman, W.E. 1981. Diversity relations and succession in Californian coastal migration corridors. of species from southern climates sage scrub. Ecology 62: 170–184. Finally, we offer a word of cau- into more northerly areas to prevent tion in applying species distribution the extinction of local plant popula- Erin C. Riordan and Philip W. Rundel, modeling to conservation and re- tions. Policymakers need to consider Department of Ecology and Evolutionary source management. Our findings the costs of avoiding massive local Biology, University of California, Los An- represent one possible future trajec- die-offs and habitat conversions as geles, Hershey Hall, 612 Charles E. Young tory of change for CSS and should part of their preparation for climate Drive East, CA 90095-7246, eriordan@ be viewed as a hypothesis of how change. The costs of adapting to cli- gmail.com; [email protected]

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 7 The west side of the Arica Mountains within the Palen/McCoy Wilderness with carpets of cinch weed (Pectis papposa). THE ARICA MOUNTAINS: BOTANICAL EXPLORATIONS OF A PHYTOGEOGRAPHICAL BLACK HOLE by Duncan S. Bell

ocated in eastern Riverside were many rare plant communities dunes. The summit elevation of the County about 30 miles west of and several hundred different spe- highest peaks is around 2,100 feet. Vidal Junction and south of cies, most of which had never be- Curious about this unexplored Highway 62, the Arica Moun- fore been reported. range, I returned just a few months Ltains are what botanists refer to as a later in March of 2009. The differ- “phytogeographic black hole” be- ence was striking. Thousands of cause there is little to no floristic FROM ROCK AND SAND flowering plants had sprung forth information from the area (see map, COME LIFE from the harsh rocky mountains that page 9). Prior to my explorations of had initially looked barren. the Arica Mountains, the only col- I first visited the Aricas in De- Being new to the world of botany, lections known from the area were a cember 2008 and found a harsh the Arica Mountains became my few roadside specimens from High- rocky mountain range nearly devoid classroom. Essential training in- way 62 to the north and along the of life except for a few near-dormant cluded how to make plant collec- eastern side of the mountain range. shrubs and cacti. The area receives tions, how to take notes on habitats What I found when I explored the little annual rainfall (variable at and plant communities, and how to area more thoroughly, however, <1-5 inches) and contains many sand identify plants. I started my explo-

8 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 LOCATION MAP OF THE ARICA MOUNTIANS. rations in the center of the range, and over the course of four years worked my way into all corners of this vast section of desert that makes up the Arica Mountains and its drain- age. Presented here are just a few discoveries from this botanic educa- tion adventure.

HABITATS AND THEIR INHABITANTS

Canyons and Ridges The center of the Arica Moun- tains is made up of rugged canyons and ridges holding many plants that somehow persist despite the heavy and continuous erosion of the range. Found here in abundance are ghost growing on our planet, and with its Desert Pavement flower (Mohavea confertiflora), para- dramatic white bark and intensely On the large alluvial fans along chute plant ( platy- purple is one of the most the east side of the Arica Mountains phylla), and notched-leaved phacelia beautiful trees of our California is an extensive “desert pavement”— (Phacelia crenulata) which during deserts. Many of the paloverdes here an unusual geological formation that good years can number in the thou- grow over 30 feet tall and often have looks something like a massive jig- sands. large quantities of fallen branches at saw puzzle put together over thou- Hiding on limestone cliff faces their bases, which provide home and sands of years. This habitat is so in the center of the Arica Mountains shelter to many desert mammals, rocky and rugged that at first it may is the yellow liguled Godding’s aster reptiles, and invertebrates. Ocotillo appear devoid of life, but on closer (Xanthisma spinulosum var. good- (Fouquieria splendens) can be found inspection one can find tough desert dingii). It is not currently on the occasionally on benches above these annuals such as the elegant lilac CNPS inventory but potentially washes. All of these plants represent sunbonnet (Langloisia setosissima) could be since it is only known to one of the northernmost sections of or the devil’s spineflower (Chori- occur in a few California mountain Desert in California. zanthe rigida), along with showy pe- ranges in the upper northeast sec- One of the more unique inhab- rennials like white ratany (Krame- tion of our Colorado Desert. Closer itants of these desert washes is the ria grayi) and the colorful harshly inspection of these cliffs and break- desert Christmas tree neck slopes reveals many hidden (Pholisma arenarium), Solar projects threaten populations of crucifixion thorn wonders neatly tucked away and also referred to as sand (Castela emoryi) and other rare species in Rice Valley. nearly unreachable by any botanist food, because the Native without climbing gear. Americans would roast and eat this plant Washes/Microphyll Woodland (Moerman, 1998). It is The Arica Mountains are popu- often found parasitizing lated with great stands of micro- the roots of burrobrush phyll woodland, a plant community (Ambrosia dumosa). Of comprised of small-leaved, subtropi- similar uniqueness and cal leguminous trees that are fairly oddity is Cooper’s tall in height and grow along desert broomrape ( washes. The representative species cooperi), also a parasitic found here include paloverde (Park- plant and also found in insonia florida), smokebush (Psoro- washes punching sky- thamnus spinosus), and ironwood ward through hard com- (Olneya tesota). Ironwood is one of pacted soils with great the densest and heaviest woods force.

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 9 spined cottontops (Echinocac- tus polycephalus). All these plants are so painful to the touch that most of the fauna in the area seem to ignore them completely. Valley Floor Over 70% of the Arica Mountains drainage runs out into Rice Valley on the east side of the range and is an area of great biodiversity. Here is a desiccated valley floor that is being overtaken by sand dunes and some patches of micro- phyll woodland. This mixing Continuous erosion requires plants to be tough to survive. Two such rugged species include desert Christmas tree (Pholisma arenarium) (left) pushing up through compacted soils in one of the many of sand and rockier soils is a washes of the Arica Mountains, and notch-leaved phacelia (Phacelia crenulata) (right) growing out preferred habitat for the rare of a limestone cliff face near the summit. pink funnel lily (Androsteph-

TABLE 1. SOME RARE PLANTS OF THE ARICA MOUNTAINS.

Species Common Name California Habitat and Distribution Rare Plant Rank (CRPR) Androstephium Pink funnel lily CRPR 2.2 Sandy/rocky places on ridgeline of range and scattered breviflorum widely in Rice Valley. Astragalus insularis Harwood’s milkvetch CRPR 2.2 On sand ramp on west side of range and scattered var. harwoodii uncommonly in sandy places of Rice Valley. Astragalus Borrego milkvetch CRPR 4.3 Loose sand and compacted sandy places in the northern lentiginosus var. section of the Arica Mountains. borreganus Castela emoryi Crucifixion thorn CRPR 2.3 Large scattered population of over 2,500 individuals in Rice Valley. Chamaesyce Abrams’ spurge CRPR 2.2 Summer annual. Found only in a few localized popula abramsiana tions in Rice Valley. Cryptantha costata Ashen forget-me-not CRPR 4.3 Sandy places from north section of the Aricas out into Rice Valley. Funastrum swallow wort CRPR 4.2 A few large populations found in washes and canyons utahense on west side of Aricas. Ditaxis claryana Glandular ditaxis CRPR 2.2 A single population found on rocky outcropping being taken over by sand dunes on west side of Aricas. Ditaxis serrata California ditaxis CRPR 3.2 A localized population found in rugged rocky canyon in var. californica center of the Arica Mountains. Eriastrum Harwood’s eriastrum CRPR 1B.2 Sandy places on west side of Aricas and in extended harwoodii dunes from north of range out into Rice Valley. Nemacaulis woolyheads CRPR 2.2 A few localized populations on sand ramp on west side denudata var. of Aricas, and a single population in Rice Valley. gracilis Proboscidea Devils claw CRPR 4.3 A few localized populations in small drainages in Rice althaeifolia Valley.

Source: Duncan Bell, 2013.

10 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 LEFT: Ghost flower (Mohavea confertiflora) growing out of rocky scree on an open ridgeline in the Arica Mountains. RIGHT: Harwoods eriastrum (Eriastrum harwoodii), one of the rare plants growing in the sand dunes of the Arica Mountains. ium breviflorum) and, after the sum- is one of the largest in the California onto rocky slopes and mountain mer monsoonal storms, devils claw deserts. The amount of life in these sides, can be found on the west side or unicorn plant (Proboscidea al- dunes is staggering. On a wet year of the mountain range. These are thaeifolia). the area is blanketed with millions similar to the other sand dunes of Of great interest and importance of flowering annuals that are cov- the range except that everything is is the large and widely scattered ered by hundreds of different inver- distributed much more vertically. In population of crucifixion thorn tebrate species. Hastening to take the center of the Arica Mountains a (Castela emoryi). With approxi- advantage of this seasonal, lush sand ramp blows ever so slightly mately 2,500 plants, this is currently green desert bounty, many fly fran- over the ridgeline onto its eastern the largest population in California tically from plant to plant collecting side, which is so steep that the sandy and the world. This large robust as much nectar and pollen as pos- sections often collapse upon them- shrub has very long, spiny branches sible, while others crawl in zigzags selves. Many of the plants that grow that are often tipped with robust or circles to gather their food. here have roots three to ten times seed clusters. If you want to see this These dunes are home to many the length of their terrestrial bodies plant in flower you will have to brave populations of rare desert plants that help to anchor the ever shifting temperatures of 110º F or more, such as Harwood’s milkvetch (As- and unsteady slopes they call home. since it only flowers in the middle of tragalus insularis var. harwoodii), summer. Because it is one of the growing at its most northwestern SEASONS only plants on the valley floor to limit here; the bristly ashen forget- flower regularly during the summer, me-not (Cryptantha costata), which The Arica Mountains have two it can often be found with a ca- prefers deep shifting sands; the distinct flowering seasons: the spring cophony of insect life from native gentle woollyhead (Nemacaulis season that follows the winter rains, wasps to intrepid climbing ants. denudata var. gracilis), which is be- and the summer season that follows coming increasingly rarer in Cali- the monsoonal storms. The spring Sand Dunes and Sand Ramps fornia due to development that has season is more consistent, and the A great many of the rare plant destroyed most of its historic popu- abundance of flowering plants var- species of the Arica Mountains can lations; and Harwood’s eriastrum ies from year to year depending on be found in its sandy habitats. The (Eriastrum harwoodii), a curious the amount of rainfall. sand dunes that blow in from the little plant that often seems to grow My first spring season in the north into Rice Valley on the east leaning to the east. Aricas was blessed with a profu- are a continuation of those blowing Sand ramps, which are inclined sion of wildflowers. Thousands of in from Cadiz Valley to the north- or vertical sand dunes that have strongly scented notch-leaved pha- west, and together this dune system formed by wind blowing sand up celias covered the rugged slopes of

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 11 The main western drainage of the Arica Mountains with its many peaks and ridgelines in the background. the mountain range. In between without a sign of summer annuals. (Allionia incarnata), at times grow- the phacelias, many other showy After exploring this range for two ing six feet long, were so abundant annuals were to be found such as years, I was under the impression that one’s feet and legs would regu- the delicate desert star ( that no summer annuals grew there, larly get entangled in them. There bellioides), purple mat (Nama demis- as I had not seen a single one. How- was also the strong scented cinch sum), Parry’s dalea (Marina parryi), ever, in the summer of 2011 during weed (Pectis papposa) numbering in a dozen different species of popcorn my third year of exploration, I went the millions, and so abundant that flower (Cryptantha), desert dande- out after a storm blew over from after several days of exploration my lion ( glabrata), and ev- , and to my amazement boots and lower pant legs were bright erywhere desert fivespot (Eremalche found what seemed like a billion yellow with its pollen. rotundifolia). That so many delicate annuals in bloom. There were liter- Summer annuals require the desert annuals can spring up from ally thousands of spiderlings (Boer- ground to be a certain temperature the rocky rubble of such a rugged havia) of three different species, and they require monsoonal rains. mountain range is a wonder indeed. brightly colored fringed amaranth That these seed banks can lie in wait In marked contrast, the summer (Amaranthus fimbriatus), and the for so many years is amazing. And flowering season happens very ir- viney prostrate caltrop (Kallstroemia when the event happens, it all hap- regularly, and decades can go by californica). The prostrate windmills pens so fast. The temperature at the

LEFT TO RIGHT: The parasitic desert broomrape () growing among shrubs in Rice Valley. • Devils claw (Proboscidea althaeifolia) in flower in Rice Valley following some heavy monsoonal storms. • The pendulous seed pods of Harwood’s milkvetch (Astragalus insularis var. harwoodii) growing in sand dunes of the Arica Mountains. • Borrego milkvetch (Astragalus lentiginosus var. borreganus) growing in compacted sandy soils in the northern part of the range. • Delicate desert fivespot (Eremalche rotundifolia) growing out of the harsh rocky slopes of the Arica Mountains. • Wooly heads (Nemacaulis denudata var. gracilis) at sunrise growing in the sand dunes of Rice Valley.

12 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 duct, and south of Highway 62 to the current boundary area to include all sandy sections, would greatly assist in the conservation of these rare plant com- munities, including a large stand of approximately 2,500 crucifixion thorn individuals. The Arica Mountains also have a very diverse fauna and include other sensitive species such as desert tor- toise, fringed toed lizard, kit fox, badger, burrowing owl, prairie fal- con, and the cheeseweed owlfly Cottontop cactus (Echinocactus polycephalus) growing on a rocky alluvial fan on the east (CNDDB and personal observation). side of the Arica Mountains. With all other common and rare spe- cies included, the Arica Mountains time of flowering is still around 100º the form of trampling and herbivory and Rice Valley comprise a very or warmer so that some of these to the native vegetation. unique, special, and fragile ecosys- annuals can germinate, grow, flower, The most recent threat to the tem that deserves greater attention. and go to fruit in just a few weeks, area is the renewable energy projects It was actually while camping in and sometimes faster. that target open flat areas of the desert the Arica Mountains that my wife where large solar fields are being found out she was pregnant for the installed. One such project that has first time. I hope that 50 years from THREATS AND already been approved will be in- now I’ll be able to take my grand- CONSERVATION NEEDS stalled at the northern section of Rice children out to this amazing moun- Valley. The plan includes digging tain range and show them the same Unfortunately the Arica Moun- wells in Rice Valley and pumping things I found when I was their age. tains and Rice Valley face many water from the ground. This could And hopefully they will be able to threats of varying degrees, from in- be detrimental to the large popula- do the same with their grandchil- vasive non-native species to devel- tion of crucifixion thorn and many dren, so that many generations go- opment pressures. Sahara mustard other plants that grow here. ing forward can also enjoy the (Brassica tournefortii) is very prob- The sand dunes of Rice Valley beauty and diversity of this special lematic and is most prolific in the and to the north of the Arica Moun- place in our California deserts. This sandy habitats, where it is crowding tains are home to many rare plant will only be possible if we formu- out our native annuals. Off highway communities that need greater pro- late and put into action an effective vehicles (OHV) continue to destroy tection. While the western section conservation plan to address threats and degrade habitat. While Rice Val- of the Arica Mountains and the to this wild area. ley was an open OHV area over a southernmost section of Rice Valley decade ago, it is now closed to ve- lie within the Palen-McCoy Wilder- REFERENCES hicles, but people are still showing ness under the jurisdiction of the up to drive on the dunes. Rice Val- Bureau of Land Management (BLM), Baldwin, B.G., et al. 2013. The Jepson ley is also open to livestock grazing, the large expansive sections in- Manual: Higher Plants of California, which creates a lot of destruction in between are open areas susceptible Second Edition, http://ucjeps.berkeley. to destruction and edu/IJM.html. development. Ex- California Natural Diversity Database (CNDDB). 2013. RareFind 4. Cali- panding the wil- fornia Department of Fish and Wild- derness bound- life, Sacramento, CA. aries north of the Moerman, D.E. 1998. Native American Arica Mountains Ethnobotany. Timber Press, Portland, to include all OR. sandy areas be- tween the moun- Duncan S. Bell, 1500 North College tain range and the Avenue, Claremont, CA 91711-3157, dbell California aque- @rsabg.org

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 13 INDUCERS, PARASITOIDS, AND INQUILINES: LIFE INSIDE THE PLANT GALL by Peter Bryant

ost people are familiar are concerned mainly with galls in- Parasitoids: Galls provide a with the round growths duced by insects. home for the inducing species, but on oak trees that we call most of them are also invaded by a oak apples. However, wide variety of other insects. After Moak apples are just one example of a INHABITANTS OF THE a gall starts to form, very often an- diverse set of related structures called GALL other type of wasp, a parasitoid (dif- galls that are induced on various fering from a parasite in that it kills plants by mites, fungi, bacteria, and Galls are convenient to study its host, whereas a parasite does small insects including wasps, saw- since they can easily be collected not) lays one or more eggs inside flies, midges, and aphids. Ron Russo and kept in containers until some- the growing gall. The resulting larva (2007) has produced a wonderfully thing emerges. But in doing this I kills and consumes the original comprehensive field guide to the galls soon found that what emerges is inhabitant of the gall, then goes of California and other western states, only sometimes the expected prog- through metamorphosis and emerges and in Fremontia (38:4/39:1, Oct. eny of the insect that induced the from the gall to propagate the para- 2010/Nov. 2011) he has described gall; in fact there are three quite sitoid species. In some cases the his discovery of even more plant galls. different types of gall inhabitants. parasitoids have evolved extraordi- Gall formation is often thought narily long ovipositors (egg-laying to reflect a defense mechanism on Inducers: Galls form when the appendages) that allow them to de- the part of the plant, but this inter- female insect (of the inducer spe- posit eggs far inside the gall and pretation does not fit the facts very cies) lays her eggs in or on the plant, close to the target which is the in- well, since in the process of “de- and the plant responds by encap- ducer larva. This is especially im- fending itself” the plant provides a sulating the developing larva or lar- portant in large galls such as oak protected, nutritious chamber that vae in a growing chamber called the apples, which can, in fact, be as big allows the growth and development gall. In some cases the gall contains as apples. David Attenborough calls of a new generation of gall inducers. only one individual of the induc- these parasitoids “burglars” and in Since the plant usually appears to ing species (it is monothalamous), his BBC documentary, “Life in the suffer no harm, the insect/plant re- whereas in other cases it contains Undergrowth,” he shows amazing lationship here fits the definition of many larvae of the inducing species video of a long ovipositor finding commensalism (one partner ben- (it is polythalamous). its way through the gall tissue to efits, the other is not harmed or Insect galls can form at various the target insect. benefited) rather than parasitism or sites on the plant: leaf, stem, bud, mutualism. But what attracts biolo- root, catkin, or acorn. Some types Inquilines: Several types of wasp, gists to galls is that they support a stay attached, whereas others fall beetle, and moth larvae, called in- fascinating micro-ecosystem that is off. Those on and stems are quilines, live inside galls induced easily accessible for study. Here we the most conspicuous. by other insects and feed mainly on

LEFT TO RIGHT: Red, bean-like “apple galls” on the leaf of arroyo willow (Salix lasiolepis) induced by the willow leaf gall sawfly Pontania pacifica. • The sawfly is named after the saw-like ovipositor that the female uses to deposits eggs. • Larva revealed in a bisected apple gall.

14 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 the nutritive tissues. Although the among the most conspicuous of GALLS ON WILD ROSE inducer larva is not the main tar- galls, and they harbor a micro- get, it sometimes also gets killed ecosystem as fascinating as it is com- California wild rose (Rosa cali- by the inquiline as plicated. The in- fornica) develops at least three kinds it feeds on the gall ducer is the willow of galls, all induced by wasps in the tissue. leaf gall sawfly family Cynipidae. The most spectacu- The situation can (Pontania pacifica), lar are the spiny leaf galls that are get much more com- named after its saw- induced by the wasp Diplolepis po- plicated with the ar- like ovipositor that lita. These are monothalamous, as rival of additional it uses to penetrate can be seen in the photo on page 16 parasitoids, leading the leaf for deposi- to the idea of a com- tion of the egg. But munity of insects the resulting galls associated with the can be occupied by gall. Some are hyper- parasitoids from sev- parasitoids, meaning eral different fami- that they are parasi- lies of wasps, and toids on parasitoids, they are also invaded and there can be by inquilines includ- parasitoids on the in- ing larvae of a moth quilines. It is com- and a weevil. monplace for a gall Another com- type to support para- mon willow gall, the sitoids from five or willow rosette gall, six families, but there looks like a cabbage are reports of much and is induced by a more complicated peculiar mechanism communities, in- quite unlike the cluding in one case growth stimulation a report of an oak that appears to be in- gall with over 75 volved in the forma- associated species tion of most other (Wiebes-Rijks and galls. Instead, the Shorthouse 1992)! In larva of the induc- well studied cases it ing species takes up has been shown that Parasitoid wasps that emerged residence in the distinct generations from willow apple galls. TOP TO shoot tip and lives (often sexually and BOTTOM: Wasps from the families: in the exact position Pteromalidae (Pteromalus sp.), asexually reproduc- Eurytomidae, Ichneumonidae normally occupied ing generations) of (Lathrostizus sp.), and Eulophi- by the dividing cells the inducing species, dae (Pnigalio sp.). necessary for shoot parasitoids of differ- elongation. This ap- ent types (some attacking each pears to prevent the shoot from other), and inquilines all arrive at growing any more in length, while specific different times during the it continues to produce shortened life of the gall (Wiebes-Rijks and but swollen leaves that pile up on Shorthouse 1992). one another to produce the cabbage- like gall. The inducer can be attacked GALLS ON ARROYO by parasitoid wasps from at least WILLOW four families, and the gall can be invaded by at least two kinds of in- One plant that is well known for quiline beetles and an inquiline TOP TO BOTTOM: The willow rosette gall. • its galls is the arroyo willow (Salix moth. These galls stay on the tree Bisected to show the inducer, the larva of lasiolepis). The red beans that de- and can often be seen as blackened the willow rosette gall midge (Rabdophaga velop in the leaves of this plant are remnants the following season. salicisbrassicoides). • The adult midge.

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 15 where one very small larva has in- morphosis in the pupal stage and are a swollen stem gall induced by duced a gall that is huge compared emerge as an adult (unless it has Diplolepis nodulosa, and a large, ter- with the inducer. The single larva in- been attacked by parasitoids!). minal, round leafy gall induced by duces a gall containing enough nu- Wild rose produces at least two Diplolepis californica. My DNA bar- trients to allow it to grow to full size, other galls, both polythalamous, in coding studies show that the three after which it will go through meta- addition to the spiny leaf one. These wasps responsible for inducing these

TOP LEFT: Intact spiny leaf gall induced by the wasp Diplolepis polita on California wild rose (Rosa californica). TOP RIGHT: Bissected spiny leaf gall with a single larva inhabiting the gall. MIDDLE LEFT: Intact swollen-stem gall, probably induced by the wasp Diplolepis nodulosa on California wild rose. MIDDLE RIGHT: Bisected swollen-stem gall with multiple inducer larvae. BOTTOM LEFT: Intact terminal leafy gall induced by the wasp Diplolepis californica on California wild rose. BOTTOM RIGHT: Bissected terminal leafy gall showing multiple inducer larvae.

16 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 three distinct gall types are close relatives of each other, much more closely related than they are to other cyni- pid wasps. This suggests that these three species are de- scended from a common an- cestor that was already a gall inducer associated with wild rose.

OAK GALLS For some reason our lo- cal scrub oak (Quercus ber- beridifolia) develops an im- pressive variety of galls, each one induced by a different species of wasp in the fam- ily Cynipidae. Some of the galls boggle the mind, with form and color that seem totally foreign to the host plant. How can one small larva completely reprogram the plant tissues?

Two of the many gall types on California scrub oak (Quercus berberidifolia). LEFT: Multiple spined GALLS ON OTHER turban galls on a single leaf. RIGHT TOP: The inducer of spined turban galls, Andricus douglasii. • RIGHT OAK SPECIES MIDDLE: Disc gall on a leaf. RIGHT BOTTOM: The inducer of disc galls, Andricus parmula.

Just as scrub oak produces a spec- ing gall, which forms on the leaves most amazing of all galls is that, for tacular variety of galls, other species of valley oak (Quercus lobata) and some unknown reason, they jump! of oak produce their own collection other oaks in Central California. As with Mexican jumping beans, the of gall types. For example, Califor- They are among the smallest galls to larva inside the gall is able to make nia live oak (Quercus agrifolia) pro- be found, measuring only about the shell click with enough energy duces a pumpkin gall and a two- 1mm across, and they are unusual that the whole structure leaves the horned gall, and canyon live oak in that they are formed in huge num- ground. The jumping gall soars (Quercus chrysolepsis) produces a bers and are dehiscent (they fall to many times its own height before two-story gall. Both species also pro- the ground rather than remaining coming in to land again. When the duce oak apples and other more com- attached). But what makes them the larva is removed from the gall it mon galls, all induced by does continue to make specific cynipid wasps. some jerky movements, Joyce Gross has compiled and these must be respon- photographs of a huge sible for causing a rapid collection of oak galls at click movement in the http;//joycegross.com/ shell. galls_ca_ oak.php. Another mystery is why these galls jump. When they fall onto pave- THE NOTORIOUS LEFT: A beaked twig gall on California scrub oak. The internal structure ment, they seem to be re- of galls can be surprisingly well organized. In this gall, the inducing sponding to the heat of JUMPING GALL larva resides in a chamber which is connected to the shell of the gall the surface where they by numerous thin fibers. RIGHT: Two larvae inside this gall were One of the most in- identified by DNA barcoding. The larger larva is the inducing cynipid, have fallen, but when they teresting galls is the jump- and the smaller one is a parasitoid from the Eupelmid family. fall into a more normal

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 17 COTTONWOOD PETIOLE GALLS

One more example of a gall must be mentioned because its inducer is so different from all the rest. The gall forms at a very specific loca- tion—the petiole at the base of the leaf on Fremont cottonwood (Popu- lus fremontii) trees—and the inducer is an aphid called Pemphigus populi- transversus. An egg produced by sexual reproduction in the spring initiates the gall, and inside the gall the fat, wingless “stem mother” (the fundatrix) grows and gives birth to a family of fatherless juveniles. These feed on the growing tissues of the gall, eventually develop wings, and fly off to colonize the roots of sec- ondary plant hosts in the mustard family including turnip and cabbage. They reproduce asexually on the sec- Galls on canyon live oak (Quercus chry- ondary hosts, where they can cause solepis). TOP AND MIDDLE LEFT: Steeple gall, intact and bisected to show the single economic losses and are known as inducer larva. BOTTOM LEFT: The inducer, cabbage root aphids. Heteroecus sanctaeclarae. BELOW: A parasi- toid eupelmid wasp (Reikosiella sp.). SEX LIFE OF GALL INSECTS A common strategy among the cynipids is an alternation between asexual and bisexual (sexually re- producing) generations, each gen- eration often being associated with a different type of gall or host plant. In most cases the galls of the asexual generation appear during late sum- mer and fall, while galls of the sexual generation can be found in spring SAGE AND COYOTE BUSH and/or early summer (Pujade-Villar GALLS et al. 2001). Two of our local spe- cies (Andricus quercuscalifornicus Galls developing on other plants and Disholcaspis plumbella) appear are induced by midges in the family to have given up on sexual repro- Cecidomyiidae. One can be found duction altogether, since only fe- on white sage (Salvia apiana) where males and no males have been found. the galls are tubular in shape and protrude from both upper and lower leaf surfaces, and another on coy- QUESTIONS FOR FUTURE ote bush (Baccharis pilularis) where RESEARCH the gall forms a rounded swelling environment, their jumping move- at the tip of a branch. Yet another Galls may not be very significant ments might help them to retreat is found on honeysuckle (Lonicera in the grand scheme of nature, but into the shelter of leaf litter rather subspicata), where it resembles a they do raise some fascinating ques- than staying exposed at the surface. small bud. tions. The most intriguing is the na-

18 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 Jumping galls, induced on valley oak( Quercus lobata) by the wasp Neuroterus saltatorius. The galls are aligned with a 1 mm scale. ture of the signal that induces gall morphology and color of many plant family and it represents an economi- formation and maintains its growth. galls. Often it appears that the insect cally serious plant disease. Whether Much is known about the hormones, has completely reprogrammed the other gall inducers also work by “ge- including auxins, cytokinins, and plant tissue to make structures— netically engineering” their hosts is gibberellins, that control growth in including discs, stars, hairy balls, perhaps the most important ques- plants. But whether the gall induc- spotted balls, and some in bright tion in gall biology, and new tech- ers are using these growth-control colors—that are quite specific for niques in molecular biology should pathways or others has been a mys- the inducing species but are com- make it possible to answer this ques- tery until recent ground-breaking pletely novel for the plant. I have tion in the near future. work by Yoshihito Suzuki and col- often heard the comment that this leagues in Tokyo. must mean that the gall inducer REFERENCES Their work indicates that, for works by introducing new DNA into willow leaf galls induced by saw- the plant cells. If so, this would be Chilton, M.D., et al. (1977). Stable in- flies, the inducing signal is, indeed, an almost unprecedented horizon- corporation of plasmid DNA into a cytokinin precursor produced by tal transfer of genetic information higher plant cells: The molecular the sawfly and injected into the plant from one species to another. basis of crown gall tumorigenesis. at the time of egg laying. Further- Horizontal gene transfer is al- Cell 11:263–71. Costacurta, A., and J. Vanderleyden. more, they showed that subsequent most, but not totally unprecedented, 1995. Synthesis of phytohormones growth of the gall is maintained by since the crown gall bacterium by plant-associated bacteria. Critical the production of an auxin by the (Agrobacterium tumefaciens) is Reviews in Microbiology 21: 1–18 larva. They also showed that in the known to transfer a small piece of Pujade-Villar, J., et al. 2001. Current growing gall tissue there was in- DNA (a plasmid) into the plant cell state of knowledge of heterogony in creased activity of genes that are where it integrates into the chromo- Cynipidae (Hymenoptera, Cynipoi- known to be regulated by auxin and somal DNA. The introduced DNA dea). Ses. Entom. ICHN-SCL 11: 87– cytokinin. So in a remarkable evolu- contains genes for the production of 107. tionary process, the gall inducer has auxin and cytokinins, and these hor- Russo, R. 2007. Field Guide to Plant found a way to hijack the plant’s mones cause the affected cells to Galls of California and other Western States. California Natural History normal growth regulatory mecha- grow rapidly and produce a gall on Guides, 91. University of California nisms. the root crown (Costacurta and Press, Berkeley, CA. More intriguing still is the ques- Vanderlevden, 1995). The crown gall Stone, G.N., and K. Schönrogge. 2003. tion of what controls the amazing bacterium affects plants in the rose The adaptive significance of insect gall morphology. Trends in Ecology and Evolution 18: 512–522. Wiebes-Rijks, A.A., and J.D. Short- house. 1992. Ecological relation- ships of insects inhabiting cynipid galls. In Biology of Insect-Induced Galls, ed. O. Rohfritsch. Oxford University Press, New York, NY. Yamaguchi, H., et al. 2012. Phytohor- mones and willow gall induction by a gall-inducing sawfly. New Phytolo- gist 196: 586–595.

Peter Bryant, School of Biological Sci- LEFT: The petiole gall induced by the aphid Pemphigus populitransversus on Fremont cottonwood (Populus fremontii). RIGHT: The easily recognizable fundatrix and her progeny ences, University of California, Irvine, CA inside the gall. 92697, [email protected]

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 19 THE NEED FOR SITE-SPECIFIC MAINTENANCE MANUALS FOR PUBLIC AND PRIVATE NATIVE PLANT LANDSCAPES by Ellen Mackey and Nancy L.C. Steele

ost of us who are gar- Several years ago Ellen Mackey, So following several impromptu deners have probably together with Bart O’Brien and Betsey discussions with landscaping work- faced this predica- Landis, coauthored the book, Care ers at different parks in 2007 and ment at one time or & Maintenance of Southern California 2008, and thinking about some land- another.M We were at a plant sale or Native Plant Gardens (hereafter re- scapes under development, we de- nursery and couldn’t resist buying ferred to as the Care & Maintenance cided to use the Care & Maintenance some of the beautiful plants we saw Manual) (O’Brien, Landis, Mackey Manual to develop customized that were not on our list. But now 2006). It is designed to assist home- manuals and checklists for new na- we’re stumped. How big will they owners and beginning maintenance tive landscapes. Our thinking was get? Where should I plant them and staff with caring for native plant land- that workers would be much more how do I care for them? scapes, and addresses these very is- likely to use checklists tailored to Now imagine that you work for sues. The Care & Maintenance the needs of the landscapes where a public agency or community Manual, however, is encyclopedic, they were working, than to refer to group and have been handed a and requires a certain level of exper- generalized information from a newly installed landscape to main- tise. We realized that many workers book. tain. It happens to be a drought involved in the day-to-day mainte- tolerant native plant garden, and nance of landscapes did not have the BACKGROUND maybe you have little experience time to read the book, even though it caring for this type of landscape. was printed in both English and Span- The Council for Watershed How do you keep things growing ish, and often lacked the background Health, which promotes sustainable and looking good? required to make good use of it. water management in the Greater Los Angeles area, has been involved in writing or editing a number of landscaping support documents over the past ten years, including the Los Angeles River Master Plan Landscap- ing Guidelines and Plant Palettes (LADPW 2004). Unique to the Guidelines is the requirement that an ecologist work with the land- scape professional to choose and place the appropriate locally native plants selected from an extensive list. (The document can be accessed at: http://dpw.lacounty.gov/wmd/ Watershed/. Then click on Major Watersheds/Los Angeles River/Mas- ter Plan/Landscaping Guidelines.) Beginning in 2006, the Council for Watershed Health offered sus- tainable landscape seminars to train landscape professionals working on public landscapes. Through these seminars we gathered feedback on what information and approaches Looking south across Old Marengo Park in Altadena, California: the first spring, showing were working and what was not. rock signage, poppies, native shrubs, and trees. All photographs by Nancy Steele unless In addition, the Council was re- otherwise noted. quired to develop manuals and con-

20 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 goals. You should provide details on to include the original plant list as the partners, design concept, and well as the 11” x 17” planting plan, eventual maintenance responsibili- followed by species-by-species in- ties. Additional information, such as formation on the sun exposure, soil, grant contracts and press releases, pruning, pests, etc. To aid landscape could be included in appendices. personnel in identifying species, we include a picture of each plant in a 2: Soils, Planting, Fertilization, and garden landscape (as opposed to a Mulch. Document how you tested wild situation) and a close-up of the the project site soil, how to plant flower. We’ve heard too many hor- and stake trees, when and how much ror stories of maintenance crews re- to fertilize, and how to apply mulch. moving the wrong plant—killing the You might want to include photo natives and leaving more familiar documentation of the plant installa- weeds in place. A dry well inlet in Old Marengo Park was tion to track the changing landscape discovered after a pile of mulch that had as plants grow, die, and are replaced. 5: Weed Management. We conduct been covering it was removed. The under- ground dry well detains and infiltrates The soils report should be included a survey in late winter for weeds stormwater that flows across the site. either here or in the appendices. within the project area, and include Photograph by Ellen Mackey. in the maintenance manual addi- 3: Watering, Irrigation, Stormwater. tional weeds that have been found duct trainings for the Elmer Avenue Other than the planting plan, no in the area and might show up later Neighborhood Retrofit, a project that documents seem to disappear more in the project site. This additional uses bioswales and an infiltration quickly than the stormwater and ir- information helps maintenance gallery under a street to collect rigation plans. We recommend that crews separate the weeds from the stormwater on-site. We were con- you include both 11" x 17" construc- expensive native plants. We prima- cerned about how to apply the in- tion documents as part of the manual. rily use Weeds of California and Other formation in the Care & Maintenance Western States, Vol. 1 & 2 (DiTomaso Manual to a residential setting. 4: Plant Maintenance. You will want and Healy 2007) for descriptions and Serendipitously, we had the oppor- tunity to test out our ideas on an- other native landscape, a new pocket park developed with funding from Metropolitan Water District of Southern California. What follows is a chapter-by- chapter description of key informa- tion that should be included in a site-specific maintenance manual so that others can develop their own. Links to custom manuals we have created appear in the sidebars ac- companying this article. We used the Care & Maintenance Manual as the primary source of information, but also referred to California Na- tive Plants for the Garden (Bornstein, Fross, and O’Brien 2005) and to nu- merous websites, including Tree-of- Life, Theodore Payne Foundation, Las Pilitas, and CNPS.

1: Introduction. We think it is help- ful to memorialize and include a his- Closeup of native wildflowers in bloom at Old Marengo Park. Pictured are California tory of the project, along with de- poppies (Eschscholzia californica), Eaton’s penstemon (Penstemon eatonii), tidy tips (Layia sign approach, project elements, and platyglossa), and elegant clarkia (Clarkia unguiculata).

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 21 recommended management tech- could harm native plants. Describe Graffiti. All these elements require niques. Be sure to obtain permission invertebrate and vertebrate pests and maintenance as well since, for ex- to reproduce photos in your manual control methods, but also define and ample, rocks in a seating wall can from the UC Regents. This chapter describe beneficial animals, such as become loose, or a water fountain should also include the weed con- lizards and insects (e.g., ladybug lar- can break. For graffiti removal you trol methods you want to be used. vae, praying mantis, and common will want to specify acceptable meth- black ground beetle). ods (e.g., steaming), and informa- 6: Pest Management. This chapter tion about who to call and whether addresses possible animal pests that 7: Hardscape, Site Amenities, and surfaces have an anti-graffiti coat-

22 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 OLD MARENGO PARK he Old Marengo Park is an T8,000-square-foot, gently sloping triangle that lies on the southwest corner of Wood- bury Road and Marengo Av- enue, bordering the cities of Altadena and Pasadena. For 40 years this parcel was a dirt and gravel unimproved eyesore. Neighbors living on the “Old Marengo” cul-de-sac have long lived with this dirt lot across from their homes. In 2007 the Arroyos & Foot- hills Conservancy, a 501(c)(3) nonprofit corporation, and the Altadena Watershed Commit- tee, a community organization, completed a waterwise oak woodland pocket park for this old county right-of-way. The oak woodland references near- by mountains and appears to pull a piece of upland habitat down to the urban Altadena/ Pasadena border. The park reflects the histori- cal and natural heritage of Altadena and Pasadena, and acts as an informal gateway be- tween the two communities. The Council, with support from Barbara Eisenstein, crafted a maintenance manual for the park in 2008. The lot size and limited planting list was the per- fect venue to develop our main- tenance manual and checklist templates. Our target audience was the volunteers’ supervisor who needed support through an individualized manual and monthly checklist (http://mm. ing. This chapter could also include responsible for removing these ar- watershedhealth.org). information on any public art that is ticles, as well as for removing litter. located on the site and the 11” x 17” hardscape plan. 9: Green Waste Management. With not be mulched because their seeds advance planning and staff training, are likely to spread infestations on 8: Litter management. Our public maintenance crews can shred green site. Describe what equipment may landscapes seem to attract throw- waste into mulch and keep it on be needed, and list contacts. away items such as couches and site. However, be sure to include a mattresses. Be sure to list who is warning about weeds that should Appendices. This is the place to in-

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 23 Monthly Checklist. Our monthly checklist consists of an 11" x 17" spreadsheet that lists all tasks, both general and species-specific, for a particular landscape. This way a crew supervisor can quickly assess which tasks are necessary for the month and allocate time accordingly. Since is also included, this eliminates any guesswork about how to take care of these native plants.

We hope that this site-specific and species-specific maintenance manual for native landscapes will serve as a template for professionals (ecologists, landscape architects, de- signers, and contractors) to create their own. Such a resource will in- crease the probability of success and Rock and gravel-lined swale on Elmer Avenue fills up with stormwater on a rainy day. the long-term beauty of these na- Note the meandering sidewalk and dymondia (Dymondia margaretae) in the 18” next to tive landscapes. We’re also inter- the curb for passengers exiting cars. The landscaped area includes Douglas iris (Iris ested in hearing from readers about douglasiana), spreading rush (Juncus patens), sanddune sedge (Carex pansa), autumn sage (Salvia greggii), and germander sage (Salvia chamaedryoides). any improvements that you discover which will make maintenance clude copies of the soils report, all might want to list nonprofit organi- manuals even more useful to land- detailed drawings of the irrigation zations (with their permission, of scape practitioners. and stormwater components, as well course) that can provide educational as the planting details for trees and information, such as the California REFERENCES shrubs. We also include emergency Native Plant Society and the Califor- Bornstein, C., D. Fross, and B. O’Brien. contacts for the police and fire de- nia Invasive Plant Council, and vol- 2005. California Native Plants for the partment, trash collectors, and local unteer groups, such as the commu- Garden. Cachuma Press. Los Olivos, graffiti removal resources. You also nity service organization Big Sunday. CA.

ELMER AVENUE NEIGHBORHOOD RETROFIT he Elmer Avenue Neighborhood Retrofit is the first complete “green street” in the City of Los Angeles, Tcreating a walkable street that conserves more water than residents use, on average. The Council for Watershed Health and its partners retrofitted a one-block section of Elmer Avenue in Sun Valley with state-of- the-art storm water best management practices to capture and infiltrate runoff, reduce flooding, improve water conservation, reduce pollution to the Los Angeles River, restore habitat, provide green space, and beautify the community. Specific techniques include bioswales and subsurface galleries for infiltration, native drought tolerant landscapes, smart irrigation controllers, permeable surfaces, and solar streetlights. In 2012 the Council completed construction on the Elmer Paseo, a mid-block permeable walkway with biowales that were planted with all natives. This neighborhood-scale project provides a real-world model of sustainable design and serves as an example of how to provide multiple benefits along with community enhancement. In addition, the Council has been monitoring Elmer Avenue before the project began and has accumulated a wealth of information on plant survival, water quality improvements, and soil accumulation. The Council gave each resident a customized maintenance manual and checklist, in Spanish and English, designed for a non-professional audience (http://mm.watershedhealth.org).

24 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 THE COUNCIL FOR WATERSHED HEALTH he Council for Watershed Health is a Thub for watershed research and analysis in Southern California. It is uniquely posi- tioned to influence and inform policy by convening forums and conducting applied research that is reliably fair, objective, and rooted in science. Founded in 1996, the Council uses an inclusive process to improve the economic, social, and ecological health of our water- sheds through applied research, education, and planning. The Council promotes Vision 2025—a model of sustainable, urban water- shed management for Southern California. This collaborative vision promotes clean wa- ter, reliable local water supplies, restored native habitats, ample parks and open spaces, integrated flood management, and revital- ized rivers and urban centers. The Council’s Sustainable Landscape Seminars offer workshops on sustainable landscape design and management prac- tices, with experts from the fields of land- scape architecture, ecological restoration, ornamental horticulture, arboriculture, and land care. Speakers share current research, best practices, and case studies, and facili- tate field sessions. To further its goals, the Council has ini- tiated the Native Seed Resources Coalition (NSRC). The goal of this stakeholder pro- cess is to nurture an ethically obtained, reliable supply of locally native plants for public landscaping and restoration projects and residential gardens. For more information on the seminars, go to: http://watershedhealth.org (Symposia & Seminars/Sustainable Landscape Semi- nars). For more information on NSRC, go to Elmer Avenue vegetated swale with maturing plants veiling the gravel- http://nativeseed.watershedhealth.org. lined swale on a sunny day. Sanddune sedge (Carex pansa), spreading rush (Juncus patens), Germander sage (Salvia chamaedryoides), moonshine yarrow (Achillea ‘Moonshine’), bearded iris (Iris sp.).

DiTomaso, J., and E. Healy. 2007. Weeds Los Angeles Department of Public nia. Metropolitan Water District of of California and Other Western States. Works. http://ladpw.org/wmd/water- Southern California, Los Angeles, Volumes 1 & 2. University of Califor- shed/LA/LAR_planting_guidelines_ CA. nia, Division of Agriculture and webversion.pdf. Natural Resources, Publication 3488. O’Brien, B., B. Landis, and E. Mackey. Ellen Mackey, 700 N. Alameda Street, Los Angeles County Department of 2006. Care & Maintenance of South- Los Angeles, CA 90012, emackey@ Public Works. 2004. Los Angeles ern California Native Plant Gardens. mwdh2o.com; Nancy L.C. Steele, 700 N. River Master Plan, Landscaping Guide- Cuidado y mantenimiento de jardines Alameda Street, Los Angeles, CA 90012, lines and Plant Palettes, ed. E. Mackey. de plantas natives del sur de Califor- [email protected]

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 25 SEQUOIA: CONTINUATION OF THE SAGA by Marcel Rejmánek

he redwood—Sequoia sem- and most likely correct: the name waterlily). Haenke’s herbarium spe- pervirens (D. Don) Endl.—is Sequoia is not a celebration of the cimens had quite complicated fates. one of the three iconic trees famous Cherokee Indian chief Haenke died under uncertain cir- of California, the other two Sequoyah, but originated as a posi- cumstances in Cochabamba, Bolivia, beingT the giant sequoia (Sequoiaden- tion of this in Endlicher’s in 1816 (Stearn 1973). dron giganteum (Lindl.) J. Bucholz) highly artificial suborder Cunning- What happened to specimens he and the bristlecone pine (Pinus hameae where Sequoia was placed was sending to Madrid is not clear. longaeva D.K. Bailey). It is not sur- in a sequence (therefore Sequoia) However, it was only in 1821 that prising that so much has been writ- among genera ordered by numbers seven chests with about 15,000 plant ten about this species. One of the of seeds per cone scale. This is an specimens, which had lain neglected last issues of Fremontia contained intriguing story. Inevitably, there are for many years in Cadiz and the port an 11-page article dedicated to this many other interesting questions we of Hamburg, were acquired by the taxon (Lowe 2012). It was rather may ask about our redwoods. Here recently funded Bohemian National entertaining, having been written are just a few of them. Museum in Prague and thus came almost like a detective story, but into the hands of Haenke’s compa- essentially about the origin of the WHO WAS THE triot botanists. Inevitably, many of redwood’s scientific name. BOTANICAL DISCOVERER the taxa that were subsequently de- Lowe’s conclusion is plausible OF THE REDWOOD? scribed as new had already been named elsewhere. Nevertheless, According to some authors names of some of the Californian (McClintock 1989, McCarthy 2008), species described on the basis of the first botanist to discover the red- Haenke’s specimens are accepted wood was British surgeon and even in the new Jepson Manual (e.g., naturalist Archibald Menzies of the Epilobium brachycarpum C. Presl, George Vancouver expedition, in Juncus falcatus E. Mey., Luzula como- 1794. According to others (Lowe sa E. Mey., Oenanthe sarmentosa C. 2012), Menzies was the first one Presl ex DC., Poa secunda J. Presl, who made the proper botanical col- Scirpus microcarpus J. & C. Presl). lection of this species. However, Willis Linn Jepson in his The Silva of California in 1910 WHAT ARE THE (p. 138) writes that the discoverer REDWOOD’S CLOSEST was Thaddeus Haenke, a botanist of RELATIVES? the Malaspina Expedition of 1791. Some specimens of Sequoia collected Obviously, to follow questions by Haenke may be in Spain, but about the first botanist to collect several of his specimens are in specimens of the Sequoia or the ori- herbaria of the National Museum gin of its scientific name are inter- and Charles University in Prague. esting and entertaining enquiries. This was the material for which But there are some botanically even Karel (Carl) Borivoj (Boriwog) Presl more interesting questions as well. (1849, p. 237) used the name Se- Sequoia is one of the 32 currently One of the specimens of the redwood quoia religiosa. recognized genera in the family Cu- collected in California by Thaddeus Haenke was a blessed botanist. pressaceae (cypress family, now in- Haenke in 1791, now in the Charles In California, he discovered the tall- cluding the former Taxodiaceae). It University Herbarium in Prague. On the est tree in the world and 10 years is generally acknowledged that the labels are three species names: Sequoia later, in Bolivian Amazonia, he was nearest relative of Sequoia semper- sempervirens Endl (on the top, obviously added later), Taxiphyllum sempervirens the first botanist who found the virens in North America is Sequoia- Presl, and Taxodium sempervirens Lamb. aquatic plant species with the larg- dendron giganteum (giant sequoia). Photograph by Jirí Hadinec. est leaves, Victoria amazonica (giant In fact, until 1939 these two species

26 FREMONTIA VOL. 41, NO.3, SEPTEMBER 2013 (9). Stebbins therefore concluded that, “if the distinguishing charac- ters are all considered to be of equal importance, Sequoia must be re- garded as closer to Metasequoia than to Sequoiadendron.” Moreover, only weighting cer- tain vegetative characters (arrange- ment of leaves and deciduous vs. evergreen character) as more impor- tant than reproductive characters could lead to the opposite viewpoint. In the same article, Stebbins dis- cussed the type of Sequoia hexapoly- ploidy (the configuration of 66 chro- Simplified phylogenetic (evolutionary) relationships among genera in the family mosomes in the meiotic metaphase) Cupressaceae (after Yang et al. 2012). Genera with asterisks are currently native to and suggested that it probably origi- California. nated via hybridization between two, and perhaps three, distinct ancestral were treated by many botanists as phyll, shape of cone scales, number species. His concluding hypothesis members of the same genus, Sequoia. of cotyledons, arrangement of leaves, was that Sequoia sempervirens origi- Then, there is the third morpho- etc.) to assess similarities and differ- nated as an allopolyploid from hy- logically similar genus, Taxodium. ences among Metasequoia, Sequoia, brids between an early Tertiary or This genus is known mostly as T. Sequoiadendron, Taxodium, and Mesozoic species of Metasequoia and distichum var. distichum (bald cy- Glyptostrobus (shui-sung, Chinese some probably extinct type of taxo- press, growing in swamps of south- swamp cypress). Based on this analy- diaceous plant. east North America) and T. mucro- sis, Metasequoia resembled Sequoia natum (Montezuma cypress or ahue- in the largest number of characters IS IT A HYBRID? hete, the national tree of Mexico). (18), differing from it in only nine. Besides the fact that this genus is The most similar genus to Sequoia- Since the publication of Stebbins’ deciduous, there are also several dif- dendron was Sequoia, but the num- seminal paper in 1948, the idea of a ferences in reproductive structures ber of differences between these two hybrid origin of Sequoia has been that make Taxodium a rather distant (10.5) was somewhat larger than that only rarely questioned (Ahuja and relative to the redwood and giant between Sequoia and Metasequoia Neale 2002). Interestingly, using sequoia. The situation became more in- teresting when a new deciduous ge- nus—Metasequoia, sometimes called dawn redwood—was discovered in China in 1941. Based on fossil records, this genus was historically widespread, including in North America (see Eocene to Pliocene maps of its distribution in western North America on p. 44 in Lowe 2013). Interestingly, it was George Ledyard Stebbins, one of the past presidents of the California Native Plant Society, who immediately asked very fundamental questions about relationships among Sequoia, Sequoiadendron, and Metasequoia (Stebbins 1948). Stebbins used 27 characters (number of ovules per megasporo-

VOL. 41, NO. 3, SEPTEMBER 2013 FREMONTIA 27 Morphology. Acta Botanica Yunnanica 10: 33–37. Lowe, G.D. 2012. Endlicher’s se- quence: The naming of the genus Sequoia. Fremontia 40:1&2: 25–35. Lowe, G.D. 2013. Geologic history of the giant Sequoia and the coast red- wood. North America Research Group (Paleontology), nargpaleo. org, Beaverton, OR. Mao, K., et al. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. PNAS 109: 7793– 98. McCarthy, J. 2008. Monkey Puzzle Man. Archibald Menzies, Plant Hunter. Whittles Publishing, Dunbeath, Scotland, UK. McClintock, E. 1989. Early explora- tion in the West. Part I. Fremontia 17:3 15–18. morphological data, Li (1988) in- dron are always closer to each other Presl, C.B. 1849. Epimeliae Botanicae. ferred that Sequoia originated by hy- than to Metasequoia (Mao et al. Amadei Haase, Pragae. bridization between some ancient 2012). This may be in agreement Stearn, W.T. 1973. An introduction to K. B. Presl’s Reliquiae Haenkeanae species of Metasequoia and Sequoia- with Lowe’s (2013) hypothesis that (1825–1835). In: Reliquiae Haen- dendron, rejecting the Stebbins’ sug- both Sequoia and Sequoiadendron keanae, ed. K.B. Presl. A. Ascher & gestion that another extinct ances- are descendants of the Eocene Se- Co., Amsterdam, pp. 1–17. tor was involved. The idea that Se- quoia affinis. The saga of Sequoia Stebbins, G.L. 1948. The Chromo- quoia originated from a recombina- continues. somes and relationships of Metase- tion between genera Metasequoia and quoia and Sequoia. Science 108 (July Sequoiadendron got some new sup- REFERENCES 30): 95–98. port in the recent molecular phylo- Yang, Z-Y., J.H. Ran, and X-Q. Wang. genetic study of Cupressaceae phy- Ahuja, M.R., and Neale, D.B. 2002. 2012. Three genome-based phylog- logeny by Yang, Ran, and Wang Origins of polyploidy in coast red- eny of Cupresaceae s.l.: Further evi- (2012). A simplified picture of wood (Sequoia sempervirens (D. Don) dence for the evolution of gymno- Endl.) and relationship of coast red- Cupressaceae molecular phylogeny sperms and southern hemisphere wood to other genera of Taxodiaceae. illustrates their conclusions. biogeography. Molecular Phylogenet- Silvae Genetica 51: 93–100. ics and Evolution 64: 452–470. Nevertheless, this is very likely Jepson, W.L. 1910. The Silva of Cali- not the end of the story. In another fornia. Memoirs of the University of recent Cupressaceae phylogeny California, Volume 2. The University Marcel Rejmánek, Department of Evolu- based on nucleotide sequences from Press, Berkeley, CA. tion and Ecology, University of Califor- plastid, mitochondrial, and nuclear Li, L. 1988. The parents of Sequoia nia, Davis, CA 95616, mrejmanek@ DNA, Sequoia and Sequoiaden- sempervirens (Taxodiaceae) based on ucdavis.edu

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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- Telos Rare Bulbs tions/Fremontia. Fremontia Editorial Advisory The most complete offering of bulbs native to the Board western USA available anywhere, our stock is Susan D’Alcamo, Jim Andre, propagated at the nursery, with seed and plants Ellen Dean, Phyllis M. Faber, from legitimate sources only. Holly Forbes, Dan Gluesenkamp, Brett Hall, Todd Keeler-Wolf, David Keil, Pam Muick, Bart Telos Rare Bulbs O’Brien, Roger Raiche, Teresa P.O. Box 1067, Ferndale, CA 95536 Sholars, Greg Suba, Dick Turner, www.telosrarebulbs.com Mike Vasey, Carol Witham

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CONTRIBUTORS FROM THE EDITOR

Duncan Bell is a field botanist for Rancho Santa Ana Botanic ecently I came across an old issue of National Geo- Garden, and spends most of his time doing floristic work in graphic (February 1999) that had been given to me by the undercollected and unexplored mountain ranges of Ra friend. On the cover are the words, “Biodiversity: Southern California. The Fragile Web.” The lead article’s teaser blurb on the table of contents page reads, “Hosting an astonishing array Peter Bryant is a research professor at the University of of living things, our hospitable planet is becoming less so California, Irvine. During most of his research career he has for many species.” studied growth control and cancer genetics using fruit flies. Biodiversity is critically important to the health of wa- Ellen Mackey is a senior ecologist with Metropolitan Water tersheds throughout California and to sustaining life in District of Southern California and the Council for Water- general, although its importance is not well understood by shed Health. She has coauthored a number of documents the general public. The conservation of rare plants has on native plants in Southern California. always been at the heart of the CNPS mission, and rare plants and plant communities add greatly to a diverse flora. Marcel Rejmánek is a professor of ecology at the University For that reason alone, CNPS’s new initiative, the Rare of California, Davis. His research and teaching focus on bio- Campaign, is a timely idea. We hope it will bring in signifi- logical invasions, regeneration of tropical forests, and plant cant funds to support and expand the invaluable work of the community ecology. CNPS Rare Plant Program. At the same time, it offers us a Erin C. Riordan is a postdoctoral researcher in the Depart- fresh opportunity to sponsor a variety of public talks and ment of Ecology and Evolutionary Biology at UCLA. Her events that explain why rare plant species are important, and PhD research modeled future land use and climate change what can be done to conserve them. CNPS chapters can play impacts on California sage scrub. a significant role in making this happen, and we hope you will involve many of your members in this worthwhile effort. Philip W. Rundel is a distinguished professor of biology in And here’s a sneak preview of what’s to come. The next the Department of Ecology and Evolutionary Biology at two issues of Fremontia will be thematic ones, and will focus UCLA. His research centers on the comparative ecology and on issues related to the richness of California’s deserts. The conservation of the world’s five Mediterranean climate re- first issue, out in January 2014, will cover the unique plants, gions. diversity, and ecology of the Mojave and Sonoran Deserts. Nancy L.C. Steele is executive director of the Council for The second issue will discuss threats to desert ecosystems, Watershed Health. Nancy co-founded the Arroyos & Foot- and what is being done to protect them or minimize those hills Conservancy in 2000, and is co-owner with her hus- impacts. band of Chaparral Mountain Honey Company. —Bob Hass

FREMONTIA (continued on inside back cover) VOL. 41, NO. 3, SEPTEMBER 2013