Crossosoma Journal of the Southern California Botanists, Inc. Volume 29, Number 1 Spring-Summer 2003 CONTENTS Lichens on rock and biological crusts enhance recruitment success of rare Dudleya species (Crassulaceae) in Southern California — Richard E. Riefner, Jr., Peter A. Bolder, Thomas W. Mulroy, and Carl Wishner 1 Three notable lichen collections and their relationship to lichen distributions in Southern California — Kerry Knudsen 35 Southern California Botanists, Inc. Source and Use of Funds — 2003 39 http://www.socalbot.org Crossosoma CROSSOSOMA (ISSN 0891-9100) is published twice a year (normally about May and November) by Southern California Botanists, Inc., a California nonprofit corporation. Subscription rate to domestic libraries and institutions is $25.00 per calendar year, or $30.00 for foreign institutions (for individual membership, see inside back cover). Back issues (Vols. 18-present) are available for $5.00 an issue or $10.00 a volume, postpaid. 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See the website for email addresses that can be used for electronic submission. Views published in CROSSOSOMA are those of the contributing author(s) and are not necessarily those of the editors, the membership of Southern California Botanists Inc., or the SCB Board of Directors, unless explicitly stated. Copyright © 2003 by Southern California Botanists, Inc. All rights reserved. Permission to reproduce items in CROSSOSOMA, in whole or in part, should be requested from the current Editor. http://www.socalbot.org Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 1 Lichens on Rock and Biological Crusts Enhance Recruitment Success of Rare Dudleya Species [Crassulaceae] in Southern California Richard E. Riefner, Jr. 5 Timbre, Rancho Santa Margarita, CA 92688 rriefner@earthlink.net Peter A. Bowler Department of Ecology and Evolutionary Biology University of California, Irvine, CA 92697-2525 Thomas W. Mulroy Science Applications International Corporation 525 Anacapa Street, Santa Barbara, CA 93101 Carl Wishner Envicom Corporation 28328 Agoura Road, Agoura Hills, CA 91301 ABSTRACT: Dudleya (Crassulaceae), a genus of about 45 taxa restricted to the southwestern United States and Baja California, Mexico, has a number of species with narrowly restricted geographic distributions, specific microhabitat requirements, and low population numbers that make them vulnerable to extinction. In southern California, these include the federally listed threatened Dudleya cymosa (Lemaire) Britton & Rose ssp. agourensis K. Nakai, D. cymosa ssp. marcescens Moran, D. cymosa ssp. ovatifolia (Britton) Moran, D. parva Rose & Davidson, D. stolonifera Moran, and D. verityi K. Nakai. Other rare species include D. blochmaniae (Eastw.) Moran ssp. blochmaniae , D. multicaulis (Rose) Moran, D. variegata (S. Watson) Moran, and D. viscida (S. Watson) Moran. Rock-inhabiting lichens serve to trap seeds and to enhance seedling survival of the rare and/or threatened Dudleya species that are restricted to steep cliff and rock habitats. Fruticose lichens, especially Niebla ceruchoides Rundel & Bowler (the pincushion lichen), are richly branched, and thereby, effectively trap minute Dudleya seeds as they are dispersed by wind and rain across rock outcrops. These bush-like lichens collect airborne soil particles and absorb moisture from fog, thereby creating a microhabitat that facilitates the establishment of Dudleya seedlings in the otherwise inhospitable conditions found on sheer rock faces. 2 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] Biological crusts that grow as a thin mantle over rock or soil may also play an important role in the establishment and survival of many Dudleya species. In particular, members of the D. cymosa complex appear to be associated with moss-dominated crusts over rock. The biological association of these cryptogams with rare Dudleya species is important in designing habitat-specific conservation plans for these rare species. Habitat assessment programs must include monitoring of these lichens, bryophytes, and crust organisms not only for their own sake, but also to assure that these habitats persist in order to sustain viable populations of these rare Dudleya species. KEYWORDS: biodiversity, biological crusts, bryophytes, conservation biology, Crassulaceae, cyanobacteria, Dudleya, lichens, liverworts, mosses, Niebla, southern California, threatened species. INTRODUCTION The conservation issues and problems associated with a rapidly expanding human population are particularly acute in southern California, where large-scale urban development is leading to a massive reduction in the state’s unique and varied biota (Sala et al. 2000). This situation presents a difficult test for those interested in protecting the natural environment: can southern Californians develop effective programs to preserve the state’s rich biotic diversity? One of the challenges is to identify “vital signs,” including measurable attributes of biodiversity, and then to develop practicable approaches to monitor them. However, biodiversity has often been a minor consideration in environmental policy, because ecological relationships are often difficult to take into account (Noss 1990). One example of the progress made in developing and implementing policy pertaining to biodiversity involves a recent California Environmental Quality Act (CEQA) court decision (Superior Court of California, County of Ventura, 1998). At issue was the significant potential adverse impact from vehicle emission pollution to the pincushion lichen, Niebla ceruchoides Rundel & Bowler, and its biological association with the federally listed threatened vascular plant species, Dudleya veriryi K. Nakai (Riefner and Bowler 1995; Figure 1). In this case, the decision served both to protect the threatened Dudleya, and to strengthen the ability of CEQA to protect a lichen-dominated microhabitat that facilitates recruitment of a threatened vascular plant. Because of the awareness created by this decision, conservation biologists in southern California are now beginning to recognize that biodiversity encompasses a wide range of organisms, and is not limited to the old “laundry lists” of vascular plants, vertebrates, or vegetation types. Accordingly, the cryptogams, which have often been excluded entirely from study, and subsequently, overlooked in restoration and management programs are now considered for conservation and study. In this paper, we demonstrate the ecological importance of preserving cryptogamic organisms, and the need to recognize them in management, restoration, and monitoring programs. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 3 THE N1EBLA -DUDLEYA BIOLOGICAL ASSOCIATION The genus Dudleya Britton & Rose (Crassulaceae) is comprised of rosette-forming succulent perennial herbs adapted to arid environments, typically inhabiting ocean bluffs, sheer cliffs, and rock outcrops (Mulroy 1976; Dodero 1995). Dudleya (commonly known as “live-forever”) attains its greatest diversity along the southern California coast, the Channel Islands, Baja California, Mexico, and islands off Baja California’s Pacific coast. Many species have restricted geographical distributions, and some are very narrow endemics with specific habitat requirements (Moran 1951; Mulroy 1976). Dudleya species are particularly successful as colonizers, as few other vascular plants can endure the inhospitable conditions of rock outcrops, weathered soils, and other nutrient-poor habitats in arid southern California. Some Dudleya species grow in thin soils of rock fissures, and other species appear to form associations with lichens and bryophytes (Mulroy 1976). As described below, lichens and bryophytes may be important in the establishment of both groups of Dudleya species. In contrast, the small geophytes, such as Dudleya blochmaniae (Eastw.) Moran ssp. blochmaniae, D. multicaulis (Rose) Moran, and D. variegata (S. Wats.) Moran, are rarely found on vertical cliffs, but are most abundant about the base of outcrops, between shrubs in open patches of soil, or in grasslands (Dodero 1995; Marchant et at. 1998). Although a number of small vascular plants germinate in moss, we are not aware of another genus of vascular plants that grows in such an intimate relationship with the cliff-inhabiting lichens of the genus Niebla as species of Dudleya (Figure 2). These cryptogams act as nurse plants, providing an organic and microhabitat foot-hold on open, sheer rock that enables Dudleya to germinate, flower, and set seed in a microhabitat that otherwise would not be available for colonization (Figure 3). Within the genus Dudleya, there are species that are consistently pulverulent or glaucous from a whitish waxy deposit on the leaves, other species whose leaves are always green, and a third group with either variable pulverulence or with both a green and a “white” morphology. Based on experiments upon the green and pulverulent morphs of Dudleya brittonii D.A. Johansen, and field observations of most other species in the genus, Mulroy (1976, 1979) proposed that glaucescence (a white or bluish-silver powdery wax) is genetically controlled, and that white or green morphologies are often habitat specific: i.e., non-glaucous plants tend to occupy deeper soils; and plants that produce glaucescence are able to survive on steep, nutrient-poor, sheer rock and cliff habitats. Ecologically significant characteristics attributed to glaucescence include high reflectance of ultraviolet light, increased protection against predation, reduced susceptibility to nutrient loss caused by leaching in fog zones, and conservation of nutrient capital and biomass that enable the rapid uptake and storage of water (Mulroy 1976, 1979). Dudleya species display genetically controlled adaptations that allow them to colonize habitats unavailable to most other plants. Dudleya species also have the ability to rapidly absorb moisture when it is available by rapid production of roots. Mulroy (personal observation) found that Dudleya species produce roots in response to high humidity with no direct contact with water. The water is stored in the succulent 4 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] leaves and spongy tissue of the caudex. Like many succulents, Dudleya species use Crassulacean Acid Metabolism (CAM); these plants are usually found in environments where water availability is restricted temporarily or seasonally, such as rock outcrops (Bartholomew 1973; Gibson 1982). This water-conserving adaptation is characterized by nocturnal opening of stomata for gas exchange and Fixation of C02 into organic acids that subsequently is released during the daylight hours for use in photosynthetic carbon Fixation. The closure of stomata reduces w'ater loss due to transpiration, thereby saving water, and enabling Dudleya species to exploit exposed habitats where moisture is scarce. Perhaps these evolutionary traits also allow Dudleya to exploit the micro-scale differences of moisture, soil, and nutrient availability provided by the cryptogams in barren habitats. Because of these ecological subtleties, knowledge of demographic characteristics, ecology, and natural history are important for the conservation of genetic diversity when assessing the biodiversity of Dudleya species. The genus Niebla (Ramalinaceae) is comprised of fruticose (shrubby or bush-like) lichens that form a conspicuous component of the vegetation within the mantime fog zone from Washington to Baja California Sur, Mexico, the Channel Islands, and islands off Baja California’s PaciFic coast (Bowler et al. 1994). Moisture availability and substrate type may be important selective factors in the distribution of Niebla species (Spjut 1996), w'hich include saxicolous (growing on rock), terricolous (growing on soil), and corticolous (growing on wood, bark, or tree branches) taxa. In Baja California Norte, Mexico, Niebla josecuervoi (Rundel & Bowder) Rundel & Bowler and allied species dominate the vegetation of open-habitat soils that rival the abundance of other lichen micro- bush ( Cladina spp. and Cladonia spp.) dominated habitats of eastern North America (Rundel et al. 1972). Many Niebla species have open patterns of branching that resemble a naked winter shrub, such as Niebla homalea (Ach.) Rundel & Bowler, and others may be cushion-like (rounded) such as Niebla ceruchoides, or cespitose (clumped or tufted) like N. combeoides (Nyl.) Rundel & Bowler. Tufts of lichens may act like a comb for dew and fog condensation (Biidel & Scheidegger 1996). Apparently, a strong relationship exists between the cushion-like growth form and the functions performed by N. ceruchoides that benefit species of Dudleya growing on cliff and rock habitats. Most Niebla species fall within two general groups: the “ceruchoid” group, taxa that are characterized by a predominately terpenoid chemistry and lack well-developed chondroid (fiber- like) strands in the thaJlus (body of the lichen); and the “homalea” group that is characterized by conspicuous chondroid strands and a more diverse chemistry that includes divaricatic, barbatic, and sekikaic acids (depsides), and hypoprotocetraric, protocetraric, and salazinic acids (depsidones) (Bowler 1981; Bowler et al. 1994). Spjut (1996) transferred species of the ceruchoid group into a new genus, Vermilacinia Spjut & Hale, which is beyond the scope of discussion for this paper. Species of the ceruchoid lineage most frequently associated with Dudleya species in southern California include Niebla ceruchoides, and occasionally, N. combeoides. Niebla homalea is the most important of the homalea group. Crossosoma 29(1), Fall- Winter 2003 [issued August 2004] 5 Riefner (1992) and Riefner and Bowler (1995) made autecological observations of Niebla ceruchoides. This saxicolous lichen forms an interesting association with the threatened cliff- dwelling Dudleya stolonifera and D. verityi on coastal bluff habitats in southern California. Niebla ceruchoides is intricately branched, and its cushion-like morphology facilitates capture of minute Dudleya seeds dispersing across rock outcrops. Seed capture by these lichens appears to enhance seedling establishment in otherwise unfavorable microhabitats such as bare rock, where conditions are not favorable to germinating plants. These small lichen bushes also collect soil particles and absorb moisture and nutrients from fog, thereby providing an organic seedbed that increases overall seedling establishment of Dudleya on these rock habitats. There may be a mutually reinforcing feedback loop whereby both the lichen and the Dudleya plants increase microhabitat soil, each providing the other with a greater opportunity for colonization. Another possible mechanism by which Niebla species may increase seedling survival of Dudleya is by providing protection from insect and slug herbivory. Lichens produce chemical compounds, such as usnic acid, which are unique in nature and have been documented as a deterrent to herbivory (Lawrey 1983; Emmerich et al. 1993). As the seedlings begin to grow among or inside the lichen bushes, the lichens also physically protect young Dudleya plants from snails and other herbivores (Figure 4). These Dudleya species may have developed a resistance to these lichen substances, which have been shown to inhibit seed germination and/or growth of certain vascular plants (Pyatt 1967; Rundel 1978; Hobbs 1985; Nishitoba et al. 1987; Frahm et al. 2000). Quantitative analysis has shown that depsides and orcinol derivatives such as barbatic acid are significant “allelochemicals,” which exhibit strong growth-inhibitory properties against seedling development in the higher plants (Nishitoba et al. 1987). Although no experimental data are available, indirect evidence suggests that lichen acids may play an important role in excluding certain vascular plants from colonizing these lichen-dominated cliff microhabitats in southern California, and should be carefully considered in future research. Thus, these cliff-dwelling lichens appear to enhance seedling recruitment of these local Dudleya endemics in several ways. Not only lichens, but mosses, liverworts, and cyanobacteria that form biological crusts on rock are an often-overlooked component of the biodiversity of cliff habitats. Crust organisms have low moisture requirements, and their ability to utilize fog and dew as water sources enables them to exist where moisture deficit limits vascular plant cover and productivity (Belnap et al. 2001). Our qualitative observations indicate that these rock-inhabiting biological crust organisms, in addition to Niebla, may contribute in an important way to the establishment and survival of local cliff-dwelling Dudleya species (Figure 5). This observation is in concurrence with Mulroy (1976) who noted the association of a cryptogamic community with Dudleya in such habitats (USFWS 1999). Dodero (1995) also provides similar observations, whereby moss growing on rock acts as a rooting medium for D. cymosa ssp. marcescens (Figure 6). These relationships point to the importance of lichens and bryophytes in sustaining the biodiversity of endemic vascular plants of these habitats, and the need for broad community-based monitoring that includes these much-neglected organisms. 6 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] HABITATS AND ASSOCIATED BIOTIC FUNCTIONS OF SOME RARE OR THREATENED DUDLEYA SPECIES Toward the development of a monitoring program for the Dudleya - cryptogamic associations in southern California, we recognize three primary attributes of ecosystems described by Franklin et al. (1981): composition, structure, and function. In the sense of a whole ecosystem, these three integrated attributes encompass and define its biodiversity (Noss 1990). Composition is characterized as the identity and variety of elements in an area and is recorded in species lists, measurements of the diversity of species, and higher taxonomic levels. Structure is the physical organization or pattern of a system, including habitat complexity within a community, and micro-distributions or broad patterns at the landscape scale. Function involves ecological and evolutionary processes, including gene flow, disturbances, colonization and demographic processes, and nutrient cycling. Compositional diversity has been the traditional focus of study, and the structural simplification and disruption of functional ecological processes have not been adequately addressed (Noss 1990). One of the most important functions that lichens and bryophytes perform for colonizing Dudleya on barren rock is enhanced seedling establishment (Mulroy 1976; Riefner and Wishner 2000). Clusters of lichens and crust organisms form safe sites or “seed nests” where Dudleya can germinate if pockets of soil are lacking. The foliose (leaf-like) lichens Flavoparmelia caperata (L.) Hale, Flavopunctelia flaventior (Stirton) Hale, and Xanthoparmelia spp., Niebla spp., and the small, tufted Cladonia species are often major components of seed nests on sheer rock. Mosses, similarly, can serve as seed traps (van Tooren 1988) and germination substrates (During and van Tooren 1990), and at some localities they are better seed traps for Dudleya species than lichens (Figures 7 & 8). Seedling establishment (positive or negative) of vascular plants in bryophyte mats has been correlated with characteristics of the “turf’ structure of the carpet, moisture availability and other microclimate conditions, light intensity and quality, seasonal patterns of germination, and to allelopathic effects (During and van Tooren 1990; van Tooren 1990; Zamfir 2000). The frequency and duration of hydration or dehydration of the moss bed, and the rapidity of water loss and water uptake by individual moss species (Norris 2003), may also play an important role in the recruitment success of Dudleya in bryophyte mats. Seeds of Dudleya species germinate readily in the wild, or in the laboratory. Germination studies conducted by the Rancho Santa Ana Botanic Garden (2000) document relatively high percentages of germination, even for rare species or narrow endemics, including: D. parva - 67%; D. blochmaniae ssp. blochmaniae - 78%; D. greenei Rose - 76%; D. nesiotica Moran - 83%; D. stolonifera - 88%; and D. verityi - 62% & 90%. These data suggest that seed viability is not a limiting factor in the population dynamics of local or threatened taxa. Rather, successful dispersal and establishment into suitable habitats with favorable microclimates, soils, moisture, and so forth, with low competition may be key to the long-term persistence of these species. Lichens and bryophytes are poikilohydric; poikilohydry is the rapid equilibration of internal water content to the external environment (Mishler 2003). The uptake of water by lichens from Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 7 non-saturated atmospheres is well known, and represents essentially the reverse of evaporation (Nash 1996a). Similarly, most bryophytes take up water and nutrients through the whole plant and do not need a root system to draw them from the soil (Gignac 2001). As a result, bryophytes can survive on very hard surfaces, such as rocks, where higher plants cannot because their roots cannot penetrate the surface. Accordingly, absorption and retention of moisture by rock-inhabiting lichen and biological crust organisms may be vital to Dudleya species occupying steep rock surfaces, especially during drought, since these species can absorb moisture from dew or mist in the absence of rainfall (Figure 9). The successful recruitment of Dudleya traskiae (Rose) Moran on Santa Barbara Island depends on a narrow range of moisture conditions for germination and seedling establishment (Clark and Halvorson 1987). Knowledge of the functional attributes of cryptogams may play an important role in enhancement and restoration projects involving Dudleya species. Rock-inhabiting cryptogams may also improve nutrient cycling, including nitrogen Fixation, on cliff and rock substrates. We have observed several cyanolichens (cyanobacteria photobiont rather than green alga) that become gelatinous (jelly-like) when wet, and occasionally cyanobacteria (blue-green algae), growing in moss over rock. Cyanobacteria, and cyanolichens, such as the genus Collema, are important components of biological soil crusts that fix atmospheric nitrogen (Isichei 1990; Johansen 1993) and contribute to net sources of total nitrogen in soil environments that are otherwise nitrogen-poor (Evans and Belnap 1999). Nitrogen released from these organisms is readily utilized by vascular plants (Belnap et al. 2001). No experimental data is available, but these nitrogen-fixing organisms likely also benefit the small vascular plants surviving in cliff and rock habitats. Cliff and rock dwelling microorganisms are exceedingly important in nature, and they may mediate numerous processes in the geochemical landscape; see Dorn (1998) for a review. Epiphytic lichens (growing on trees and shrubs) of the forest communities along the Pacific Coast fog belt trap nutrients from atmospheric aerosols and fog, which otherwise would not be intercepted and utilized by the ecosystem. Munger et al. (1983) report that fog contains greater concentrations of nutrients than precipitation, by orders of magnitude. Most nutrients captured by tree-canopy lichens represent new inputs, and cycling of these minerals may be of ecosystem-level importance (Knops et al. 1991). For example, the lace lichen ( Ramalina menziesii Taylor) contributes substantially to mineral cycling and biomass turnover in the blue oak ( Quercus douglasii Hook. & Am.) savanna of California (Boucher and Nash 1990). Evidence is now mounting that lichens play important roles in rapid mineral cycling, such as the capture of allogenic nutrients that would otherwise not be retained in the ecosystem, which represent a highly significant input to nutrient- poor ecosystems (Nash 1996b). These critical ecosystem processes are unrecognized by many California biologists. Epiphytic lichens and mosses have also been functionally linked to other critical ecosystem processes, including primary production (McCune 1993). Many mechanisms that facilitate these functions have been documented, but the rapid absorption of moisture and particulates associated with aerosols appears to be one of the most important. Lichens are known to rapidly absorb moisture equal to 150%- 1200% of their dry weight (Hawksworth and Hill 1984). There is a lack of pertinent experimental data in southern California, but the importance of the biological functions of 8 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] lichens documented in forest and woodland habitats in terms of nutrient and moisture capture and recycling may be transferable to cliff and rock communities (Figure 10), which are comparatively poorly studied. Biological crusts on soil have only recently been recognized as important components of terrestrial ecosystems, playing a critical role in the functional ecology of arid lands (Eldridge 2000). These organisms likely improve the survival of several Dudleya species restricted to open- habitat soils, including species of the D. abramsii complex, D. blochmaniae ssp. blochmaniae, D. multicaulis, and D variegata. Numerous studies suggest that biological soil crusts stabilize soil surfaces and reduce erosion, improve percolation and soil moisture storage, enhance vascular plant seedling establishment, and improve soil fertility by nitrogen fixation (Belnap et al. 2001). Biological soil crusts likely benefit these and other Dudleya species in similar ways (Figure 11). HABITAT CATEGORIES McCune (1993) identified functional groups comprised of epiphytes sharing similar ecological roles and moiphology. Similarly, the distribution of saxicolous organisms is influenced by environmental characteristics: rock substrate type (volcanic, granite, sandstone), slope, aspect, and degree of shading (north-facing, shaded versus full sun), frequency of fire, associated vascular plant habitat (woodland versus scrub), and microclimate variables that include light, temperature, and moisture. The morphological groups that provide important functions to the saxicolous Dudleya - cryptogamic community habitats are the fruticose and foliose lichens, cyanolichens, cyanobacteria, the cushion or turf-forming (acrocarpous) and the highly branched mat-forming mosses (pleurocarpous), and non-seed-bearing vascular plants such as the ferns and spike-mosses. A strong relationship exists between the morphology of these organisms and the way in which they function. Accordingly, morphological groups have been proposed as surrogates for individual species in monitoring programs (Rosentreter et al. 2001). These morphological groups, and recognition of their observed and potential associated functions, could aid in a rapid assessment (Eldridge and Rosentreter 1999) of Dudleya habitats, especially on cliffs that are difficult to get to reach without disturbing sensitive Dudleya species. Table 1 provides a summary of these morphological groups, their associated functional attributes, and representative species, which are generally easy to identify that often occur in sensitive Dudleya habitats. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 9 Table 1: Description of morphological groups and associated functional attributes of the cryptogams and lower vascular plants of Dudleya habitats. MORPHOLOGICAL GROUPS HYPOTHESIZED PRIMARY FUNCTIONAL ATTRIBUTES REPRESENTATIVE SPECIES Lichens Fruticose (bush-like) Seed trapping, absorption and retention of moisture and nutrients, soil accumulation Cladonia fimbriata, C. furcata , Niebla ceruchoides , N. combeoides, N. homalea Foliose (leaf-like) Absorption and retention of moisture and nutrients, seed trapping, soil accumulation Flavoparmelia caperata, Flavopunctelia Jlaventior, Physcia dubia, Xanthoparmelia coloradoensis , X. mexicana Crustose (granular) & Squamulose (scaly) On Rock: Moisture absorption and retention Acarospora bullata, Caloplaca bolancia, Lecanora muralis, Lecidea atrobrunnea, Pertusaria flavicunda, Tephromela atra OnSoil: Soil stabilization, moisture absorption and retention, seed trapping Acarospora scheicheri, Diploschistes scruposus, Placidium squamulosum , Psora califomica, P. decipens Mosses Turf-forming (acrocarpous) Soil accumulation, absorption and retention of moisture and nutrients, germination substrate Bryum argenteum, Didymodon vinealis , Grimmia laevigata, Tortula brevipes 10 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] Mat-forming (pleurocarpous) Soil accumulation, absorption and retention of moisture and nutrients, germination substrate Anacolia menziesii, Pterogonium gracile, Scleropodium cespitans Liverworts Moisture absorption and retention, soil stabilization Asterella califomica, Riccia nigrella, R. trichocarpa Cyanolichens (gelatinous) On Rock: Nitrogen-fixation, moisture retention, soil accumulation Collema fuscovirens, Leptogium comiculatum On Soil: Nitrogen-fixation, moisture retention, soil stabilization Collema tenax, Peltula patella! a Cyanobacteria (blue-green algae) Nitrogen-fixation, moisture retention, soil stabilization Microcoleus vaginal us, Nostoc sp., Oscillatoria sp. Non-Seed BearingVascular Plants Spike-Moss (mat-like, cushion-like or cespitose) Soil accumulation and stabilization Selaginella bigelovii, S. cinerascens Ferns (erect to spreading habit with creeping underground rhizome) Soil accumulation and stabilization Cheilanthes covillei, Pellaea andromedifolia Polypodium californicum It is useful to group the habitats of the rare and threatened Dudleya species of southern California’s cismontane habitats into three broad categories, in order to provide a general characterization of the composition, structure, and functional biodiversity of their associated biological crusts and non- seed-bearing vascular plant components. These artificial categories are intended only to provide a preliminary profile of the composition and structure of Dudleya habitats, since the micro- distribution of these organisms can vary greatly with aspect, substrate, and elevation, and only a small fraction of these areas has been carefully studied. The categories are: fog-swept cliff and rock habitats; inland, lightly-shaded to exposed cliff and rock habitats; and undisturbed, open-habitat soils free of exotic weeds. FOG-SWEPT CLIFF AND ROCK HABITATS Fruticose lichens of the genus Niebla frequently dominate exposed, fog-swept cliff and rock outcrops. These lichens resemble “miniature forests” on cliff and rock outcrops along the coast of California and Baja California, Mexico. Other important components include biological crusts of mosses, foliose lichens, and tufts of smaller fruticose lichens such as Cladonia species. Crustose lichens (those appearing to be “painted” on the rock surface or appearing granular) and squamulose Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 11 forms (scaly) are almost always present. The prostrate, mat-forming pleurocarpous mosses and the short or tall turf-forming acrocarpous species are important germination substrates on locally shaded to exposed coastal cliff sites. Small ferns with long-creeping rhizomes, such as Polypodium califomicum Kaulf., and the bushy spike-moss (S. bigelovii L. Underw.), contribute to soil- building processes in rock crevices and stabilize thin soil pockets on outcrops in these habitats. Dudleya stolonifera and D. verityi are federally listed threatened species, and D. greenei is an uncommon species restricted to several of the Channel Islands. These species occupy cliff and rock habitats in the fog belt. We have not studied other rare Dudleya species that are restricted to the Channel Islands. Reference sites for these Dudleya species include mostly north-facing outcrops near Avalon on Santa Catalina Island, Los Angeles County; in Aliso Canyon and the mouth of Aliso Creek in Laguna Beach in Orange County; and Conejo Mountain, Point Mugu, and Long Grade Canyon in Ventura County. We have also examined the cliff and coastal bluff cryptogamic associations of more common but local Dudleya species at a variety of locations, including D. caespitosa (Haw.) Britton and Rose, at Abalone Point in Orange County; Motto Bay and Morro Rock Reserve in San Luis Obispo County; and D.farinosa (Lindley) Britton and Rose at Big Sur and Point Lobos in Monterey County. Species associated with these specialized environments include: Mosses: Didymodon vinealis (Brid) Zander, Grimmia laevigata (Brid.) BricL, Homalothecium nevadense (Lesq.) Ren. & Card, and Tortula brevipes (Lesq.) Broth. Liverwort: Asterella califomica (Hampe) Underw. Lichens: Acarospora bullata Anzi, Buellia halonia (Ach.) Tuck., B. stellulata (Taylor) Mudd, Caloplaca bolancia (Tuck.) Herre, C. rosei Hasse, Cladonia furcata (Hudson) Schrader, Collema crispum (Hudson) F.H. Wigg., Flavoparmelia caperata, Flavopunctelia Jlaventior, Lecania dudleyi Herre, Lecanographa hypothallina (Zahlbr.) Egea & Torrente, Lecanora gangaleoides Nyl., L. rupicola (L.) Zahlbr., Lecidella carpathica Korber, Leprocaulon microscopicum (Vill.) Gams ex D. Hawksw., Neofuscelia verruculifera (Nyl.) Essl., Niebla ceruchoides , N. combe aide a, N. homalea, Pertusaria flavicunda Tuck., Physcia dubia (Hoffm.) Lettau, P. tribacia (Ach.) Nyl., Physconnia spp., Rinodina bolanderi H. Magn., Tephromela atra (Hudson) Hafellner, Umbilicaria phaea Tuck., and several species of Xanthoparmelia. Spike-Moss: S. bigelovii. Fern: Polypodium califomicum. Lichens proposed for rare status in southern California collected from fog-zone outcrops associated with sensitive Dudleya species include: Gyalecta herrei Vezda, Phaeophyscia kairamoi (Vainio) Moberg, Punctelia punctilio (Hale) Krog, and Roccella fimbriata Darbish (Magney 1999). Uncommon lichens of coastal sensitive Dudleya habitats include Caloplaca stantonii W.A. Weber ex Arup, Dimelaena califomica (Magnusson) Sheard, Lecanora xanthosora B.D. Ryan & Poelt, N. isidiascens Bowler, Marsh, T. Nash, & Riefner, and N. robusta (R. H. Howe) Rundel & Bowler. Indeed, these habitats themselves are very limited, and there are relatively few sites with the complete community well represented. These sites should be protected. As with the lichens, the conservation of bryophytes is long overdue. Continuing studies on the 12 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] bryoflora of California may find that these south coast cliff and rock outcrop communities also support rare species of mosses and liverworts. INLAND LIGHTLY-SHADED TO EXPOSED CLIFF AND ROCK HABITATS Outside of the coastal fog belt, Dudleya species of cliff and rock habitats are most often found growing with biological crusts dominated by mat- and turf-forming mosses, liverworts, foliose lichens, and the black cyanolichens, such as species of Leptogium. Our general observations concur with those of Nash et al. (1979) who state that lichen abundance and species richness decrease with increasing distance from the Pacific Ocean; importantly, the saxicolous fruticose lichens are directly associated with the maritime microclimate influence, and disappear inland. As with the fog-belt association, the structural complexity within a community, its micro-distribution within a cliff system, or the broader pattern found at the regional scale vary greatly, and are beyond the scope of this paper. In addition to a rich bryoflora, these outcrops support several species of small ferns that are important soil consolidators. Also, the bushy spike-mosss is common in rock crevices and many open habitats on soil throughout cismontane southern California (Wilken 1993). This inland habitat supports several local Dudleya species, including the federally listed threatened D. stolonifera , D. cymosa ssp. agourensis, D. cymosa ssp. ovatifolia D. cymosa ssp. marcescens, and the rare D. viscida. Reference sites for these habitats include north-facing or shaded outcrops: near Cotharin Road, Little Sycamore Canyon of the western Santa Monica Mountains in Los Angeles County, Agoura Hills area of the central Santa Monica Mountains in Los Angeles County; Laguna Canyon, Saddleback Peak, and San Juan Canyon in Orange County; San Marcos Creek in San Diego County. We have also examined the inland cliff and outcrop cryptogams associated with several broad-ranging Dudleya species, including D. cymosa ssp. pumila (Rose) K. Nakai, D. edulis (Nutt.) Moran, and D. lanceolata (Nutt.) Britton & Rose, in the upper Fremont Canyon region, Cleveland National Forest, Orange County; the San Jacinto Wilderness in Riverside County; at Colby Canyon in San Bernardino County; at Julian, the Cuyamaca Mountains, and on Palomar Mountain in San Diego County. Xanthoparmelia mougeotii (Schaerer) Hale is an uncommon lichen (Magney 1999) that co-occurs with Dudleya viscida on granitic outcrops at several inland sites. Dudleya stolonifera is apparently unique since it grows along the immediate coast within the fog belt, and at somewhat inland sites beyond the maritime influence, mostly on north-facing outcrops. In north Laguna Canyon, which does not support large fruticose lichens or members of the obligate fog-zone cryptogamic community, mat- and turf-forming mosses support several hundred plants of this threatened live-forever. At other inland sites, such as Montclef Ridge in Ventura County, small ferns and spike-moss stabilize the soil of rock crevices that provides Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 13 habitat for the threatened D. parva. We have not studied the cryptogams and lower vascular plants associated with other rare Dudleya species found in the central portion of the state. Species associated with these specialized environments include: Mosses: Anacolia menziesii (Turn.) Par., Brachythecium sp., Bryum capillare Hedw., Didymodon vinealis , Homalolhecium nevadense , Pterogonium gracile (Hedw.) Sm., Scleropodium cespitans (C. Muell.) L. Koch, and Syntrichia princeps (DeNot.) Mitt. Liverwort: Asterella califomica. Lichens: Acarospora socialis H. Magn., Aspicilia cinerea (L.) Korber, Caloplaca modesta (Zahlbr.) Fink, Cladonia fimbriata (L.) Fr., C. pyxidata (L.) Hoffm., Collema fuscovirens (With.) J.R. Laundon, Dermatocarpon reticulatum H. Magn., Dimelaena radiata (Tuck.) Hale & Culb., Flavoparmelia caperata, Lecanora muralis (Schreber) Rabenh., Lecidea atrobrunnea (Lam. & DC.) Schaerer, Leptogium corniculatum (Hoffm.) Minks, Phaeophyscia spp., Physcia clementei (Sm.) Lynge, Thelomma mammosum (Hepp) A. Massal., Umbilicaria phaea Tuck., and several species of Xanihoparmelia, including X. coloradoensis (Gyelnik) Hale, X. cumberlandia (Gyelnik) Hale, and X. mexicana (Gyelnik) Hale. On shaded or north-facing cliff and rock outcrops with recurrent saturation or seepage. Cyanobacteria such as Microcoleus vaginatus (Vauch) Gom., and green algae may be present. Spike-Moss: Selaginella bigelovii. Ferns: Cheilanthes covillei Maxon, Notholaena califomica D. Eaton, Pellaea andromedifolia (Kaulf.) Fee, and Polypodium califomicum. Uncommon lichens co-occur with sensitive Dudleya species (D. viscida) on granitic outcrops at inland sites, including Lecanora mellea W.A. Weber. OPEN-HABITAT SOILS Undisturbed stabilized open-habitat soils are the bare, to sparsely vegetated microhabitats generally associated with nutrient-poor substrates. They occur in many plant communities, and over a wide range of geomorphic surfaces in southern California. Characteristically, these soils support a conspicuous and diverse biological soil crust comprised commonly of turf-forming mosses, squamulose and crustose lichens, fruticose lichen species ( Cladonia ), cyanolichens of the genus Collema, and cyanobacteria, which mostly grow within a few millimeters of the soil surface, and occasionally, the leaf-like Xanihoparmelia lichens. The bushy spike-moss is common, and the mesa spike-moss, Selaginella cinerascens Maxon, is often intimately associated with rare Dudleya species on mesas in San Diego County and Baja California, Mexico. The most notable feature of encrusted open-habitat soils is the near absence of exotic weeds, regardless of soil type or plant community (Riefner and St. John 2000). Species associated with this specialized environment include: Mosses: Bryum argenteum Hedw., B. capillare, Desmatodon convolutus (Brid.) Grout, Didymodon vinealis, Homalolhecium nevadense, Scleropodium cespitans, Timmiella crassinervis (Hampe) L. Koch, and Tortula brevipes. Liverworts: Asterella califomica, Riccia nigrella DC., R. sorocarpa 14 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] Bisch., and R. trichocarpa M.A. Howe. Lichens: Acarospora schleicheri (Ach.) A. Massal., Amandinea punctata (Hoffm.) Coppins & Scheid., Candelariella vitellina (Hoffm.) Miill. Arg., Cladonia pyxidata, Collema tenax (Sw.) Ach., Diploschistes scruposus (Schreber) Norman, Peltula patellata (Bagl.) Swinscow and Krog, Placidium lacinulatum (Ach.) Breuss, P. squamulosum (Ach.) Breuss, Psora califomica Timdal, P. decipiens (Hedwig) Hoffm., Psora nipponica (Zahlbr.) Gotth. Schneider, Toninia spp., Trapeliopsis califomica McCune & Camacho, and occasionally Xanthoparmelia coloradoensis and X. cumberlandia. Cyanobacteria: species of Microcoleus, Nostoc, and Oscillatoria. Spike-Mosses: Selaginella bigelovii, S. cinerascens. Open soil habitat supports several local Dudleya species: D. blochmaniae, D. multicaulis, D. variegata, and members of the D. abramsii complex. Reference sites for these species include many localities in the foothills of the Santa Ana Mountains, and San Clemente State Beach in Orange County; the Cuyamaca Mountains, Del Mar, La Jolla Canyon, Mission Trail Park, Otay Mountain, San Onoffe State Beach, and Torrey Pines Reserve in San Diego County; the “pebble plains” in San Bernardino County; and Long Grade Canyon and Conejo Mountain in Ventura County. Dudleya brevifolia (Moran) Moran, which is endemic to the concretionary soils of the Lindavista Formation in San Diego County (Cochrane 1985), is one of the species that almost always does not grow in close association with well-developed biological soil crusts. Uncommon or rare lichens associated with Dudleya species and open-habitat soils in southern California include Caloplaca cf. subpyraceella (Nyl. in Hasse) Zahlbr., Cladonia thiersii S. Hammer, Solenopsora “ cladonioides" B.D. Ryan & Timdal (in prep., see Nash et al. 2002), Texosporium sancti-jacobi (Tuck.) Nadv., and Trapeliopsis steppica McCune & Camacho (Riefner et al. 1995a; Magney 1999; Nash et al. 2002; McCune et al. 2002). Two special-status liverworts, Geothallus tuberosus Campb. and Sphaerocarpos drewei Wigglesw. (Tibor 2001) share similar habitat requirements with rare geophytes in San Diego County, and may co-occur with D. variegata. FUNCTIONAL ATTRIBUTES OF CRYPTOGAMS AND MONITORING OF DUDLEYA HABITATS Monitoring for sustained biodiversity is practical when causes and effects, probabilities, interactions, and alternative hypotheses are taken into account by a program that addresses current management programs and policies (Noss 1990). A monitoring program should document trends in seedling establishment and population dynamics of rare and threatened Dudleya species. To do this we propose selecting several species from each morphological-functional group as indicators of ecosystem health. Such a program could examine the following: (1) air pollution and fire impacts to lichens and other organisms that function as germination substrates or “seed nests;” (2) species which enhance nutrient inputs to the ecosystem such as nitrogen fixing cyanobacteria and cyanolichens; and (3) the “guild” of species that promote moisture retention and soil-building processes. Brief discussion of a few examples that could be included in a monitoring program, which may also be important in habitat enhancement efforts follow: Crossosoma 29(1 ), Fall-Winter 2003 [issued August 2004] 15 long-term information regarding trends in population dynamics. Ozone is formed from the interaction, mediated by sunlight, between nitrogen oxides and unbumed hydrocarbons produced by vehicle exhaust (Richardson 1992), and has become an important atmospheric pollutant in southern California and elsewhere as vehicular traffic increased dramatically following World War II. For lichens, oxides of nitrogen are one of the most harmful components of air pollution in the Los Angeles basin Figure 1. The pincushion lichen, Niebla ceruchoides, is intricately branched and its cushion- like morphology facilitates capture of minute Dudleya verity’ i seeds dispersing across rock outcrops in the western Santa Monica Mountains, Ventura County. These small lichen bushes also collect soil particles and absorb moisture and nutrients from fog, which provides an organic seedbed that increases overall seedling establishment of this federally-listed threatened plant species in otherwise unfavorable microhabitats such as bare rock. 16 Crossosoma 29( 1 ), Fall-Winter 2003 [issued August 2004] Figure 2. Dudleya caespitosa often grows among the densely branched mats Niebla homalea on high sea cliffs at Abalone Point in Orange County and in the central coast, including Morro Rock in San Luis Obispo County, which is shown in this photograph. The miniature forest of lichens provides an organic and microhabitat foothold on open, sheer rock that enables Dudleya to germinate, flower, and set seed where other annual or perennial vascular plants cannot survive. Crossosoma 29( 1 ), Fall-Winter 2003 [issued August 2004] 17 Figure 3. Niebla ceruchoides also forms associations with the threatened cliff-dwelling Dudleya stolonifera on coastal bluff habitats in Aliso Canyon, Orange County. Seed capture by these lichen cushions enhances seedling recruitment across expanses of barren rock, where conditions are not favorable to germinating plants. There may be a mutually reinforcing feedback loop whereby both the lichen and the Dudleya plants increase microhabitat soil and moisture, each providing the other with a greater opportunity for establishment in a microhabitat that otherwise would not be available for colonization. 18 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] Figure 4. Native landsnails. Helminthoglypta tudiculata, often congregate among outcrops for shelter during the daylight hours. Niebla and other lichens may increase seedling survival of Diidleya by providing protection from herbivory. including Dudleya verityi shown here at Long Grade Canyon. Ventura County. Lichens produce chemical compounds, such as usnic acid, which act as a deterrent to grazing by snails and slugs. Lichen chemicals are also significant alleochemicals. which exhibit strong growth-inhibitory properties against seedling development in many higher plants. Crossosoma 29( 1 ). Fall-Winter 2003 [issued August 2004] 19 Figure 5. Lichens, mosses, liverworts, and cyanobacteria form biological crusts on rock that are an often-overlooked component of the biodiversity of cliff habitats. Crust organisms have low moisture requirements, and their ability to utilize fog and dew as water sources enables them to exist where moisture deficit limits most vascular plants from establishment. Our qualitative observations indicate that these rock-inhabiting biological crust organisms, in addition to the larger fruticose Niebla, may contribute to the establishment and survival of local cliff-dwelling Dudleya species, including Dudleya stolonifera shown here in north Laguna Canyon, Orange County. 20 Crossosoma 29( 1 ). Fall-Winter 2003 [issued August 2004] Figure 6. Mat-forming species of moss ( Pterogonium gracile ) growing on rock act as a rooting medium for D. cymosa spp. marcescens on steep cliffs in the western Santa Monica Mountains. Los Angeles County. These relationships point to the importance of both lichens and bryophytes in sustaining the biodiversity of endemic vascular plants of cliff and rock outcrop habitats. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 2 Figure 7. Members of the Dudleya cymosa complex appear to be associated with moss-dominated crusts over rock. The Agoura Live-Forever, shown here, is established in moss ( Anatolia menziesii), which traps seeds that fall from above, on breccia outcrop crevices in the Agoura Hills, Los Angeles County. Crossosoma 29(1 ). Fall-Winter 2003 [issued August 2004] 22 Figure 8. Bryophyte crusts that grow as a thin mantle over sheer rock play an important role in the establishment and survival of many Dudleya species on vertical rock faces. Poikilohydry and desiccation tolerance (the ability to recover after being air-dry at the cellular level) defines many aspects of bryophyte ecology, which allows them to grow on habitats that rooted plants cannot. Dudleya species display genetically controlled adaptations that allow them to colonize these moss-dominated microhabitats. Depicted here is Anacolia menziesii in Laguna Beach, which provides a germination substrate for the threatened Dudleya stolonifera on otherwise barren rock surfaces. Crossosoma 29(1). Fall-Winter 2003 [issued August 2004] 23 Figure 9. Absorption and retention of moisture by rock-inhabiting lichens and moss crusts may be vital to Dudleya species occupying steep rock surfaces, especially during drought, since these species can absorb moisture from dew or mist in the absence of rainfall. In Laguna Beach. Dudleya stolonifera has the ability by rapid production of roots to absorb the moisture captured and retained by moss (Bryum sp.) growing on sheer rock. The water is stored in the succulent leaves and spongy tissue of the caudex. 24 Crossosoma 29( 1 ). Fall-Winter 2003 [issued August 2004] Figure 10. The rapid absorption of moisture and particulates associated with aerosols appears to an important function of the pincushion lichens growing on rock outcrops, which directly benefit the recruitment of Dudleya species in these barren, soil-free habitats. The importance of the biological functions of lichens documented in forest and woodland habitats in terms of nutrient and moisture capture may be transferable to these cliff and rock communities. Crossosowa 29( 1 ), Fall-Winter 2003 [issued August 2004] 25 Figure 1 1. Biological soil crusts stabilize soil surfaces and reduce erosion, improve percolation and soil moisture storage, enhance vascular plant seedling establishment, and improve soil fertility by nitrogen fixation. Biological crusts dominated by the lichens Diploschistes scruposus (white) and Placidium squamulosum (brown) enhance seedling recruitment and survival of Dudleya multicaulis on clayey soils at this site in the Santa Ana Mountains, Orange County. 26 Crossosoma 29( 1). Fall-Winter 2003 [issued August 2004] Figure 12. Impacts to populations of Dudleya verityi occurred following the Green Meadows fire of 1993 in Ventura County. Re-colonization of this Dudleya on several charred outcrops occurred after seed nests consisting of regenerating foliose (Xanthoparmelia [green] and Physcia [gray]) and fruticose ( Niebla ) lichens were re-established, approximately between 1999-2000. The pinholes and grooves or cryptogamic imprints provide a micro-foothold for lichen re-establishment on barren rock, which leads to further colonization by the Dudleya. Crossosoma 29( 1 ), Fall-Winter 2003 [issued August 2004] 27 (Boonpragob et al. 1989). Acidic gaseous forms of nitrogen (HNO„ HNO,) are also likely important in southern California as well (T.H. Nash III, pers. communication). See the following publications dealing with lichens and air pollution in southern California for additional reference (Nash and Sigal 1980; Sigal and Nash 1983; Blum et al. 1989; Boonpragob and Nash 1991; Nash and Sigal 1998). In the summer-dry Mediterranean climate of southern California, dry deposition is the primary pathway for accumulation of these pollutants in the lichens (Boonpragob and Nash 1990). However, van Dobben and ter Braak (1999) documented that not all lichens can be used as general pollution indicators, since some species may not be sensitive to all types of pollutants. Pollution index values that reflect variations in traffic densities and lichen species sensitivities would be expected to correlate specific degrees of urbanization with air pollution (Levin and Pignata 1995), and should be developed in southern California if we are to understand the distribution of many lichen species. Individual species of bryophytes also differ in response to air pollution, especially to mineral ions of chromium, nickel, copper or sulfur (Norris 2003), but may not be as vulnerable to damage from nitrogen oxides. Accordingly, the mosses could be expected to play a greater role in seedling recruitment of Dudleya on rock and cliff habitats if the lichens become threatened by air pollution. However, like the canary in the mine, bryophytes can alert us to very subtle changes at very local scales (Shevock 2001), such as changes in moisture regimes owing to habitat alterations. Fire is devastating to soil crust organisms (Johansen et al. 1984), and damage to saxicolous crusts has been poorly studied (Garty 1992). Hebert and Meyer (1984) observed that saxicolous crustose lichens can survive rangeland fire, and die only if touched by flame or sparks. Little is known about the post-fire re-colonization of epiphytic lichens in the southwest (Romagni and Gries 1997). Management programs that incorporate frequent prescribed bums could impact late-successional epiphytic lichens (Bowler and Riefner 2000), and these practices could also damage cliff and rock cryptogamic communities. However, no quantitative data are available regarding the response of epiphytic lichens, rock-inhabiting biological crusts, or soil crust organisms to fire in southern California. Accordingly, an assessment of fire damage to these organisms and associated post-fire re-colonization should be incorporated into a long-term monitoring program. Our general observations suggest that the fruticose lichens are most easily damaged, and the crustose lichens and turf forming mosses the least, as fire sweeps over outcrops. Severe impacts to some populations of Dudleya verityi occurred following the Green Meadows fire of 1993 in Ventura County. Re- colonization of Dudleya on several charred outcrops occurred after seed nests consisting of foliose and fruticose lichens were re-established, approximately between 1999-2000 (Riefner, pers. observation; Figure 12). The increased severity of fire in southern California scrub habitats has likely contributed to the destruction of many rock and soil crust communities, which have not been studied. Damn et al. (1983) and Garty (1992) introduced the concept of the “cryptogamic footprint,” which is the micro-relief on rock surfaces (pinhead holes and microgrooves) created by living lithobiontic microorganisms (rock-inhabiting unicellular green algae, cyanobacteria, free-living micro-fungi, and endolithic lichens) during pre-fire periods. These imprints trap and accumulate moisture, minute particles of ash and soil, and organic materials that provide a micro-site foothold that stimulate post- 28 Crossosoma 29( 1 ), Fall-Winter 2003 [issued August 2004] fire recovery of new communities of cryptogams on the same substrates. Natural plant communities usually develop an extensive underground network of mycorrhizal fungi that interconnect root systems. This network of roots and hyphae perform and mediate numerous important ecosystem functions for their host plants, including enhanced water and nutrient uptake and enhanced seedling development. Little work has been completed regarding the mycorrhizas of Dudleya. In southern California, only Dudleya greenei has been found to be a host for mycorrhizal fungi (Koske and Halvorson 1989), but few Dudleya species have been tested. Recent investigations at the pebble plains in the San Bernardino Mountains have found that D. abramsii Rose ssp. affinis K. Nakai, which grows in open-habitat soils densely permeated with mycorrhizal hyphae (Riefner and St. John 2000), is not a host. Mycorrhizal relationships of the threatened cliff-dwelling Dudleya species, especially those associated with moss crusts, have not been examined. Parke and Linderman (1980) believe that moss species may promote the survival of vesicular- arbuscular mycorrhizal fungi (VAMF) during periods that are unsuitable for host plant growth. Therefore, mosses may provide for the maintenance of an adequate supply of VAMF inoculum to enable vascular plant colonization of inhospitable habitats. This moss/mycorrhizal relationship could be very important for the rare and threatened Dudleya species associated with bryophytes. Woodward et al. (1999) propose incorporating symbiotic associates such as mycorrhizal fungi into monitoring programs, because of their importance to their hosts, which represents a collective effort to obtain resources. Identifying this relationship, however, may be problematic in Dudleya. Certain Dudleya species studied by Mulroy (1976) and Dodero (1995) produce “rain roots” for rapid uptake of water. These presumably drought-deciduous fine roots may be in a decomposed condition, or unsuitable for mycorrhizal evaluation at most times of the year, and therefore, may require a special protocol for identification (St. John, pers. communication). Woodward et al. ( 1999) also propose incorporating other measures such as identifying keystone species (organisms that have strong interactive effects with other species) in such a program. Niebla ceruchoides may be a keystone to enhanced seedling establishment of some Dudleya species on fog-swept cliffs, and similarly, the mosses may be key to recruitment success of Dudleya species on inland shaded cliffs and outcrops. Qualitative and quantitative research has identified that other cryptogams are potential keystone species: the reindeer lichen. Cladonia rangiformis Hoffm. of lowland heath communities in the United Kingdom (Newsham et al. 1995), the cyanobacterium. Microcoleus vaginatus , in southern California vernal pools (Riefner and Pryor 1996), and the moss genus Sphagnum in Canadian peatlands (Rochefort 2000). There is a serious gap in our knowledge of the life histories of the lower plants and lichens of southern California, and other essential relationships between cryptogamic species and vascular plants may be discovered. Unfortunately, there is not a list of specific traits or a protocol to identify keystone species. Accordingly, selecting measures of resilience, persistence, process, and organization is a logical starting point at which to begin a long-term monitoring program (Woodward et al. 1999) of habitats supporting rare Dudlyea species. Crossosoma 29( I ), Fall-Winter 2003 [issued August 2004] 29 Further experiments could be designed that would indicate if these Dudleya species have developed a resistance to specific lichen substances known to inhibit vascular plant growth, which could help explain their association with the lichen micro-bushes and other properties of the substratum (John W. Thomson, pers. communication). Finally, quantitative data are needed to document the frequency with which Dudleya germinates and sprouts within Niebla thalli and bryophyte mats in natural conditions. CONCLUSION Currently, experimental data are severely limited with regards to air pollution effects of individual lichen and moss species, keystone species associations, damage to crust organisms by changes in the fire ecology, and other factors that could impact populations of rare Dudleya species in southern California. Establishment of baseline conditions is needed while viable populations of these lichen, bryophyte, and Dudleya associations are still available for study. Lichens, mosses, fungi, and the algae have not been adequately taken into account by preservation efforts in southern California. Although they are essential to healthy ecosystems, academic botanists, consulting biologists, and land managers have largely overlooked these organisms; they clearly deserve greater attention by resource agencies (Faber 1998; Shevock 1998). Development and implementation of a monitoring program for the Dudleya - cryptogamic associations in southern California could address the goals proposed by Eldridge (2000) by creating a broad-based awareness of cryptogams, their habitats, and their benefits to biodiversity. ACKNOWLEDGMENTS John W. Thomson, University of Wisconsin, Madison, reviewed an early draft of this manuscript and helped with identification of lichens collected during the study. Chicita Culberson, Duke University, James D. Lawrey, George Mason University, Reed F. Noss, Conservation Science, Incorporated, Fred Roberts, F.M. Roberts Publications, Roger Rosentreter, Bureau of Land Management, Ted St. John, BioNet LLC, and Darrell Wright, California Lichen Society (CALS), also reviewed a draft and provided helpful comments. We also thank Thomas H. Nash III for reviewing the section on air pollution, and Theodore L. Esslinger for conducting a literature search regarding lichen acids and allelopathy. Thomas Nash and Kerry Knudsen (CALS) shared information compiled during the Sonoran Desert Lichen Project. Michael Wall, Rancho Santa Ana Botanic Garden, provided information on Dudleya seed germination, and John Tiszler, National Park Service, Santa Monica Mountains National Recreation Area, provided information on the Green Meadows fire. We are also grateful to several resource ecologists for granting permits to study the cryptogams in California state parks. COPYRIGHT RESTRICTIONS 30 Crossosoma 29(1 ). Fall-Winter 2003 [issued August 2004] The photographs in this article may not be reproduced in any format without written permission of the first author. LITERATURE CITED Bartholomew, B. 1973. Drought response in the gas exchange of Dudleva farinosa (Crassulaceae) grown under natural conditions. Photosynthetica 7:1 14-120. Belnap. J.. B. Biidel. and O.L. Lange. Definition of biological soil crusts. In: Belnap. J. and O.L. Lange (eds.). Biological soil crusts: structure, function, and management. Spnnger-Verlag Berlin Heidelberg. Belnap. J.. J.H. Kaltenecker. R. Rosentreter. S. Leonard. J. Williams, and D. Eldridge. 2001. Biological soil crusts: ecology and management. United States Department of the Interior. Bureau of Land Management, and the United States Geological Survey; Technical Reference 1730-2. Blum. O.B.. T.H. Nash III. and R. Gebauer. 1989. The effect of nitrate on CO. exchange in the epiphytic lichens Ramalina menziesii Tayl. and Pseudocyphellaria anthraspis (Ach.) Magn. from central California, pp. 181-185. In: Noble. R.D.. J.L. Martin and K.F. Jensen (eds.). Air Pollution Effects on Vegetation. Northeastern Forest Experiment Station: Broomall. PA. Boonpragob. K... T.H. Nash III. and C.A. Fox. 1989. Seasonal deposition patterns of acidic ions and ammonium to the lichen Ramalina menziesii Tayl. in southern California. Environmental and Experimental Botany 29:187-197. Boonpragob. K.. and T.H. Nash III. 1990. Seasonal variation of elemental status in the lichen Ramalina menziesii Tayl. from two sites in southern California: evidence for dry' deposition accumulation. Environmental and Experimental Botany 30:415^128. Boonpragob. K.. and T.H. Nash III. 1991 . Physiological responses of the lichen Ramalina menziesii Tayl. to the Los Angeles urban environment. Envir. Exp. Bot. 31:229-238. Boucher. V.L., and T.H. Nash III. 1990. The role of the fruticose lichen Ramalina menziesii in the annual turnover of biomass and macronutrients in a blue oak woodland. Botanical Gazette 151:114-118. Bowler. P.A. 1981. Cortical diversity in the Ramalinaceae. Can. J. Bot. 59: 437-452. Bowler. P.A.. R.E. Riefner. Jr.. P.W. Rundel. J. Marsh, and T.H. Nash III. 1994. New species of Niebla (Ramalinaceae) from western North America. Phytologia 77:23-37. Bowler. P.A.. and R.E. Riefner. Jr. 2000. Prescribed burning impacts on late successional species, pp. 71-73. In: Keeley. J.E., M. Baer-Keeley. and C.J. Fotheringham (eds.). Interface Between Ecology and Land Development in California. U.S. Geological Survey Open-File Report 00-62. Biidel. B.. and C. Scheidegger. 1996. Thallus morphology and anatomy, pp. 37-64. In: Nash. T.H. Ill (ed.). Lichen biology. Cambridge University Press. Clark. R.A.. and W.L. Halvorson. 1987. The recovery of the Santa Barbara live-forever. Fremontia 14:3-6. Cochrane. S. 1985. Dudleya brevifolia: California's own "living stone." Fremontia 13:21. Danin. A.. R. Gerson. and J. Garty. 1983. Weathering patterns of hard limestone and dolomite bv endolithic lichens and cyanobacteria: supporting evidence for eolian origin of Terra-Rossa soil. Crossosoma 29(1), Fall- Winter 2003 [issued August 2004] 31 The photographs in this article may not be reproduced in any format without written permission of the first author. LITERATURE CITED Bartholomew, B. 1973. Drought response in the gas exchange of Dudleya farinosa (Crassulaceae) grown under natural conditions. Photosynthetica 7:1 14-120. Belnap, J., B. Biidel, and O.L. Lange. Definition of biological soil crusts. In: Belnap, J. and O.L. Lange (eds.). Biological soil crusts: structure, function, and management. Springer-Verlag Berlin Heidelberg. Belnap, J., J.H. Kaltenecker, R. Rosentreter, S. Leonard, J. Williams, and D. Eldridge. 2001. Biological soil crusts: ecology and management. United States Department of the Interior, Bureau of Land Management, and the United States Geological Survey; Technical Reference 1730-2. Blum, O.B., T.H. Nash III, and R. Gebauer. 1989. The effect of nitrate on CO2 exchange in the epiphytic lichens Ramalina menziesii Tayl. and Pseudocyphellaria anthraspis (Ach.) Magn. from central California, pp. 181-185. In: Noble, R.D., J.L. Martin and K.F. Jensen (eds.). Air Pollution Effects on Vegetation. Northeastern Forest Experiment Station: Broomall, PA. Boonpragob, K., T.H. Nash III, and C.A. Fox. 1989. Seasonal deposition patterns of acidic ions and ammonium to the lichen Ramalina menziesii Tayl. in southern California. Environmental and Experimental Botany 29:187-197. Boonpragob, K., and T.H. Nash III. 1990. Seasonal variation of elemental status in the lichen Ramalina menziesii Tayl. from two sites in southern California: evidence for dry deposition accumulation. Environmental and Experimental Botany 30:415-428. Boonpragob, K., and T.H. Nash III. 1991. Physiological responses of the lichen Ramalina menziesii Tayl. to the Los Angeles urban environment. Envir. Exp. Bot. 3 1 :229-238. Boucher, V.L., and T.H. Nash III. 1990. The role of the fruticose lichen Ramalina menziesii in the annual turnover of biomass and macronutrients in a blue oak woodland. Botanical Gazette 151:114-118. Bowler, P. A. 1981. Cortical diversity in the Ramalinaceae. Can. J. Bot. 59: 437-452. Bowler, P.A., R.E. Riefner, Jr., P.W. Rundel, J. Marsh, and T.H. Nash III. 1994. New species of Niebla (Ramalinaceae) from western North America. Phytologia 77:23-37. Bowler, P.A., and R.E. Riefner, Jr. 2000. Prescribed burning impacts on late successional species, pp. 71-73. In: Keeley, J.E., M. Baer-Keeley, and C.J. Fotheringham (eds.). 2nd Interface Between Ecology and Land Development in California. U.S. Geological Survey Open-File Report 00-62. Biidel, B., and C. Scheidegger. 1996. Thallus morphology and anatomy, pp. 37-64. In: Nash, T.H. Ill (ed.). 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The Jepson manual: higher plants of California. University of California Press, Berkeley. Woodward, A., K.J. Jenkins, and E.G. Schreiner. 1999. The role of ecological theory in long-term ecological monitoring: report on a workshop. Natural Areas J. 19:223-233. Zamfir, M. 2000. Effects of bryophytes and lichens on seedling emergence of alvar plants: 36 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] evidence from greenhouse experiments. Oikos 88:603-61 1. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 37 Three Notable Lichen Collections AND THEIR RELATIONSHIP TO LICHEN DISTRIBUTIONS in Southern California Kerry Knudsen 33512 Hidden Hollow Drive Wildomar, California 92595 kzz999 @ netzero.com ABSTRACT: The model of lichen distribution in Southern California emerging from the Sonoran Lichen Flora Project is a mosaic with many missing pieces. Three collections of rare lichens from Southern California are discussed here: Catapyrenium squamellum (Nyl.) J.W. Thomson, Stromaiella bermudana (Riddle) Henssen, and Texosporium sancti-jacobi (Tuck.) Nadv. The significance and questions these collections raise in relation to our knowledge of lichen distributions in Southern California are discussed. KEYWORDS: Ascomycetes, lichen, Caliciaceae, Lichinaceae, Southern California lichen floristics, Catapyrenium squamellum, biological crust, San Mateo Canyon, Stromatella bermudana, cyanolichens, calciphiles, San Jacinto Mountains, Texosporium sancti-jacobi, Santa Monica Mountains, Agoura Hills, Verrucariaceae. INTRODUCTION Our knowledge of lichen distribution in Southern California is currently being summed up in the three volumes of the Lichen Flora of the Greater Sonoran Desert Region (Vol. 1, 2002; Vol. 2, 2004; Vol. 3 in ed.) which includes Imperial, Orange, Riverside, Santa Barbara, San Diego, and Ventura Counties, as well as Los Angeles and San Bernardino Counties (excluding the Mojave Desert). By any scientific definition, only a part of Southern California is Sonoran. The inclusion of our flora reflects two facts. The first is our natural connection with Baja California, Mexico, which is extensively covered in the flora. The second is the historical contribution of Dr. Hermann Hasse to southwestern lichenology. He published The Lichen Flora of Southern California (1913), and many smaller articles contributing to our knowledge of lichen biodiversity. Based on the Sonoran Lichen Flora, the model of lichen distribution in Southern California emerges as a mosaic with many pieces missing. Clearly, much of the limited knowledge we have of our region is based on lichenologists collecting within their local areas, such as Charis Bratt and Shirley Tucker who have collected extensively in Santa Barbara County. Or Herman Hasse, who collected in the Santa Monica Mountains for two and a half decades, but traveled elsewhere only on scattered field trips. Many collectors have explored the maritime flora, with Rick Riefner Jr.’s work being the most outstanding example (WIS, UCI, UCR). Bruce Ryan undertook landmark surveys in the San Gabriel Mountains, and the Agua Tibia Wilderness. Numerous 38 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] collectors visited the Channel Islands, probably making their florae the best explored in Southern California. Unfortunately, the rest of the records represent the work of many collectors skimming through other areas of Southern California, often just collecting along the road. In this context, I will report three notable lichen collections that I have recently made, and discuss their relationship to our current knowledge of lichen distribution in Southern California. THREE NOTABLE COLLECTIONS Catapyrenium squamellum (Nyl.) J.W. Thomson. USA: California: Riverside County: San Mateo Wilderness Area: San Mateo Canyon; along San Mateo Trail. 33° 31.720' N, 117° 24.570' W. Elevation: 387 meters. U.S.G.S. Sitton Peak 7.5’ Quadrangle. With a sunny northeastern exposure, Catapyrenium squamellum grows in a vertical biological soil crust 2x5 meters over granite mixed with Placidium lacinulatum (Ach.) Breuss, sorediate Cladonia squamules, Diploschistes species, and mosses. After the long drought of the preceding year, the crust was beginning to crack and peel. In April, 2003, a rain storm washed the crust completely off the rock and down the slope into San Mateo Canyon. Kerry Knudsen #56 (ASU), March 18, 2003. Determined and verified by Othmar Breuss, Fall, 2003. Significance: Catapyrenium squamellum (Nyl.) J.W. Thomson is a rare lichen with a West Indies disjunctive occurrence. Its type locality is in the Santa Monica Mountains, where it was collected by Herman Hasse in March, 1897, growing with moss over rock. Othmar Breuss has seen two Hasse collections, and one is listed as near the Soldier’s Home (Breuss, pers. comm.) This area is now above Wilshire Boulevard and fully developed. It has only been collected twice since in the United States: in Trinity County, California by Dr. Harry Thiers (Thomson, 1987), and in Santa Cruz County, Arizona (Bruess, 2002 and pers. comm.). A single, disjunctive collection was made by H.F. Fox in 1991, on the island of Jamaica at 450 meters elevation on soil over limestone (Breuss, 1999). The San Mateo Canyon collection represents only the sixth collection in over a hundred years. Stromatella bermudana (Riddle) Henssen. USA: California: Riverside County: San Bernardino National Forest: San Jacinto Mountains: East Canyon at end of Morris Ranch Road: on south- east-facing slope of limestone and shale rubble among mixed chaparral above Cedar Springs Trail 33° 39.407’N 116° 35.099’W. Elevation: 1734 meters. U.S.G.S. Palm View Peak 7.5’ Quadrangle. Stromatella bermudana was collected growing in cracks of limestone. Kerry Knudsen #472 (UCR) and Kate Kramer, September 17, 2003. Significance: Stromatella bermudana, a cyanolichen, is a strict calciphile. It occurs in Bermuda, its type locality. Previously, it has been collected twice in disjunctive locations in the southwestern United States, once in central Arizona, and once in Southern California (Schultz 2002). This represents its third collection in US and second time in Southern California. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 39 Texosporium sancti-jacobi (Tuck.) Nadv. USA: California: Los Angeles County: Santa Monica Mountains: Agoura Hills: un-named ridge above Cornell Comers. 34° 08.406’N 118° 45.312’W. Elevation: 375 meters. Texosporium sancti-jacobi was found at four locations on ridge (see label of voucher for other coordinates) growing on soil, Selaginella bigelovii Underw., and on the thallus of a terricolous Rinodina species, next to boulders of Conejo volcanics on a thin- soiled ridge. Kerry Knudsen #598 (ASU) and Tarja Sagar Nov. 1 1, 2003. Significance: This collection continues the discovery of new Texosporium sancti-jacobi sites in Southern California. The lichen remains rare. CONCLUSION The lack of extensive collections of lichens in Southern California makes it impossible to form hypotheses about the distribution of lichens in this region that are capable of any deductive tests. Unlike our vascular flora, the data is simply not available. For instance, is Catapyrenium squamellum really that rare? How common is Stromatella bermudana on other Southern California limestone formations? How many more locations of Texosporium sancti-jacobi could we find? Fundamental questions like these have to be answered first before we can advance to the far more intriguing questions of species distributions including phytogeography and macroecology. Answers to such questions would lead to informed decisions about the management of our biodiversity. When it is completed, the Sonoran Flora Project will give Southern California botanists an increased opportunity to develop accurate distributional data of its lichen biodiversity, as well as to discover un-described species. The three volumes of the Sonoran flora will not only give us up-to-date taxonomic revisions of all known species of our lichen flora, but also a baseline model of lichen distribution. This is a rare and historic opportunity. It is one we should build on. ACKNOWLEDGMENTS Special thanks to Dr. Othmar Breuss of the Naturhistorisches Museum, Wein, Austria, Dr. Kate Kramer and Mary Thomas of the US Forest Service, John Tiszler and Tarja Sagar of National Park Service, and Carl Wishner. Thanks to James Lendemer for reviewing the manuscript. 40 Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] LITERATURE CITED Breuss, Othmar. 2002. " Catapyrenium " Lichen Flora of the Greater Sonoran Desert Region, Vol. 1, 2002., pgs. 125-128. Breuss, Othmar. "On some pyrenocarpous lichens from the West Indies" Linzer Biologische Beitrage 31/2 31.12.1999 pgs. 839-844. Nash III, Thomas H., Bruce D Ryan, Corinna Gries, and Frank Bungartz. 2004. Lichen Flora of the Greater Sonoran Desert Region. Vol. 1, 2002. Vol.2, 2004. Tempe, Arizona: Lichens Unlimited, Dept, of Plant Biology, Arizona State University. Available by mail: http://ces.asu.edu/ASULichens/sonoran/flora.html Schultz, M. 2002. "Stromatella" Lichen Flora of the Greater Sonoran Desert Region, Vol. 1, 2002, pgs. 475-476. Thomson, John. “The lichen genera Catapyrenium and Placidiopsis in North America. ” The Bryologist: 90(l):27-39. Crossosoma 29(1), Fall-Winter 2003 [issued August 2004] 41 Southern California Botanists, Inc. Source and Use of Funds - 2003 Bank Balances at December 31. 2002: Certified Deposit Account 37,201.10 Money Market Account 5,256.28 Checking Account 8.051.55 Total 50,508.93 (Outstanding Checks 2002) -00.00 50,508.93 Receipts for Year: Membership Dues 3,671.94 Publication Sales 498.66 T-Shirt Sales 90.00 Symposium 1,729.00 Interest Income 972.84 Donations 303.05 Sales Tax 00.00 Shipping and Handling 0.00 Petty Cash 150.00 Other Income 00.00 Total Receipts 7.415.49 Total A vailable 5 7, 924. 42 Expenses for Year: Mailing 450.00 Printing 2,022.15 Postage 111.00 Symposium 692.25 T-Shirt Sales 00.00 Grants 1,000.00 Office Supplies 69.72 Typing 0.00 2002 Sales Tax 114.00 Petty Cash 150.00 Donations 00.00 Web Site 60.00 Publications 49.75 Miscellaneous Expenses 30.00 Total Expenses -4.748.87 42 Crossosoma 29(1), Fall- Winter 2003 [issued August 2004] 53,175.55 Balance at December 31, 2003 Bank Balances at December 3 1 , 2003 Certified Deposit Account Money Market Account Checking Account Total 43,124.57 3,290.12 7.290.86 53,705.55 (Outstanding checks 2002:#1631 $10.00, #1633 $20.00, #1642 $500.00) -530.00 Ending Balance 53,175.55 — Alan P. Romspert, Treasurer, Southern California Botanists Southern California Botanists, Inc. — Founded 1927 — Memberships, Subscriptions, Back Issues Individual and Family Memberships in SCB are $15 per calendar year domestic, and $20 per year to foreign addresses. Membership includes two issues of CROSSOSOMA , and 5 or 6 issues of Leaflets, the newsletter of SCB. Leaflets provides time-dated information on activities and events that may be of interest to our membership. A subscription to CROSSOSOMA is available to libraries and institutions at the domestic rate of $25 per calendar year, and $30 to foreign institutions. Back issues (Volumes 18-present) are available for $5 each, or $10 for the volume, postpaid. Prior to Volume 18, CROSSOSOMA included time-dated notices to the membership and was published six times a year. These back issues of Volumes 1-17 are $1 each, or $6 per volume, postpaid. Some back issues that are out of stock may be provided as photocopies. SCB Special Publications No. 1 A Flora of the Santa Rosa Plateau, by Earl W. Lathrop and Robert F. Thome, 39 pp $7.00 No. 3 Endangered Plant Communities of Southern California, Proceedings of the 15th Annual SCB Symposium, edited by Allan A. Schoenherr, 1 14 pp $12.00 [Special Publication No. 2, Flora of the Santa Monica Mountains, 2nd ed., by Peter H. Raven, Henry J. Thompson, and Barry A. Prigge is out of print] Book prices include California state sales tax, handling, and domestic postage. By request, the following article has been reprinted as a separate, with covers, and is available for plant collecting workshops: Reprint. Herbarium Specimens as Documents: Purposes and General Collecting Techniques, by T. S. 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Volume 29, Number 2 Fall-Winter 2003 CONTENTS The vascular flora of the Owens Peak eastern watershed: A review of progress [Sierra Nevada, Kern Co., California]. — Naomi Fraga 41 Photograph: Challenging terrain of the Owens Peak eastern watershed — Carl Wishner 44 The Vascular Plant Checklist for the University of California Natural Reserve System's San Joaquin Freshwater Marsh Reserve. — Peter A. Bowler, and Mark A. Elvin 45 Malacothrix saxatilis (Nutt.) Torrey & A. Gray var. saxatilis (Asteraceae) discovered in Orange County, Southern California — Richard E. Riefner, Jr., and Steve Boyd 67 Book Review: Coast Redwood: A Natural and Cultural History by Michael Barbour, Sandy Lydon, Mark Borchert, Marjorie Popper, Valerie Whitworth, and John Evarts (2001) — Allan A. Schoenherr (Reviewer) 70 Index to and Bibliography Volume 29, 2003 72 http://www.socalbot.org Crossosoma CROSSOSOMA (ISSN 0891-9100) is published twice a year (normally about May and November) by Southern California Botanists, Inc., a California nonprofit corporation. Subscription rate to domestic libraries and institutions is $25.00 per calendar year, or $30.00 for foreign institutions (for individual membership, see inside back cover). Back issues (Vols. 18-present) are available for $5.00 an issue or $10.00 a volume, postpaid. Prior to Volume 18, CROSSOSOMA was published six times a year; these back issues are $1.00 each, or $6.00 per volume, postpaid. SCB BOARD OF DIRECTORS FOR 2003 President Sandy Leatherman (2003) First Vice President Robert F. Thome (2003) Second Vice President Scott White (2003) Secretary Susan Hobbs (2003-2004) Treasurer Alan P. Romspert (2003-2004) Directors-at-large Terry Daubert (2003-2004) Kate Kramer (2002-2003) James Harrison (2002-2003) Tasha LaDoux (2002-2003) William Jones (2002-2003) Susan Schenk (2003-2004) Allan A. Schoenherr (2003-2004) Valerie Soza (2002-2003) Gary Wallace (2003-2004) Ex officio Board Members Sandy Leatherman (Immediate Past President, 2002) Naomi Fraga (Webmaster) Steve Leonelli (Editor of Leaflets) Carl Wishner (Editor of CROSSOSOMA ) Applications for membership, or requests for subscriptions or back issues should be sent to: Alan Romspert, Treasurer, Southern California Botanists, Department of Biology, California State University, Fullerton, California 92834, U.S.A. Notices of a time-dated nature (field trips, workshops, symposia, etc.) to be included in the newsletter Leaflets should be submitted to Micaele Maddison, the current 2004 Editor of Leaflets , do LSA Associates, Inc. 1 Park Plaza, Suite 500, Irvine, California, 92614-5981 USA. Articles, book reviews, or other items for submission to CROSSOSOMA should be sent to Carl Wishner, Editor of CROSSOSOMA , at 5169 Dumont Place, Woodland Hills, California, 91364-2309 USA. See the website for email addresses that can be used for electronic submission. Views published in CROSSOSOMA are those of the contributing author(s) and are not necessarily those of the editors, the membership of Southern California Botanists Inc., or the SCB Board of Directors, unless explicitly stated. Copyright © 2003 by Southern California Botanists, Inc. All rights reserved. Permission to reproduce items in CROSSOSOMA, in whole or in part, should be requested from the current Editor. http://www.socalbot.org CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 41 THE VASCULAR FLORA OF THE OWENS PEAK EASTERN WATERSHED: A REVIEW OF PROGRESS Naomi Fraga Rancho Santa Ana Botanic Garden 1500 N. College Avenue Claremont, CA 91711 naomi.fraga@cgu.edu ABSTRACT: Owens Peak and its eastern watershed lie at the southern end of the Sierra Nevada, within the BLM Owens Peak Wilderness Area in Kern County, California. Owens Peak itself is the highest in the Southern Sierra rising to more than 2,600 m. The location's floristic composition is unusual, possessing elements of the southern Sierra Nevada, the Great Basin, and the Mojave Desert. The flora of this area has been poorly documented, and a systematic inventory is needed to catalogue the location’s diverse plant life. Owens Peak and its eastern watershed is known to support several rare taxa including seven endemic to the Southern Sierra and two taxa discovered and described since 1984. The aim of this project is to produce a comprehensive flora of the Owens Peak eastern watershed, and to provide a foundation upon which other botanical work may be conducted in the Owens Peak area. KEYWORDS: Eastern Sierra, endemic, flora, floristics, inventory, Kern County, Mojave Desert, phytogeography, Owens Peak, rare taxa. Sierra Nevada. INTRODUCTION Owens Peak and its eastern watershed lie at the southern end of the Sierra Nevada Range, within the Owens Peak Wilderness Area (Bureau of Land Management) in Kern County, California. Elevations within the study site range from 2,600 m (8,400 ft) to 800 m (2,600 ft). Owens Peak itself is the highest point in the Southern Sierra, with the crest rarely reaching above 6,000 ft south of Owens Peak and Walker Pass (Twisselman 1967). The prominent locations within the study site include three canyons that make up the watershed and three peaks that define the ridgelines that enclose the canyons (Grapevine Canyon, Short Canyon and Indian Wells Canyon; Owens Peak, Mount Jenkins and Morris Peak from north to south respectively). The Owens Peak eastern watershed is a 50 square mile area containing enormous botanical and topographic diversity. The region’s geographic location makes for unique plant associations, as it exhibits floristic influences of the Southern Sierra Nevada, the Great Basin, and the Mojave Desert. An area of this size and diversity is interesting, but when coupled with the fact that botanical exploration has been scarce it makes such a location not only a region of botanical interest, but also a region of conservation importance. Despite the lack of systematic exploration and documentation, this area has yielded several interesting botanical discoveries in recent years, including new species and significant range extensions (Shevock 1988). The region is currently known to be the home of seven rare taxa, many of which are endemic to the Southern Sierra. These include Carlquistia muirii (A. Gray) B.G. Baldwin, Deinandra mohavensis (D.D. Keck) B.G. Baldwin, Erigeron aequifolius H.M. Hall, Eriogonum breedlovei (J.T. Howell) Reveal var. shevockii J.T. Howell, Lomatium shevockii R.L. Hartm. & Constance, 42 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Monardella beneolens Shevock, Ertter & Jokerst, Phacelia nashiana Jepson, P novenmillensis Munz, and Trifolium macilentum E. Greene var. dedeckerae (J.M. Gillett) Bameby. RESULTS AND DISCUSSION As mentioned earlier, despite having been little systematic botanical exploration in the area, work prior to this study has resulted in several interesting discoveries. These include two newly described species (since 1984); Lomatium shevockii (Hartman and Constance 1988), and Monardella beneolens (Shevock et al. 1989), and three range extensions including Deinandra mohavensis, Quercus palmeri Engelm., and Saltugilia latimeri T.L. Weese & L.A. Johnson. Deinandra mohavensis is a plant previously thought to be extinct, but recently it was rediscovered in Riverside and San Diego Counties (Sanders et al. 1997). Subsequently, it has been found within the Owens Peak eastern watershed, and further south near Cross Mountain in the Jawbone Canyon area (Steve Boyd, personal communication). This was not only a range extension for the species, but also a new record for Kem County. Saltugilia latimeri has been newly recorded at Owens Peak as a result of this study. This past field season has proved to be an excellent year for collecting, with an abundance of annuals blooming this spring. I spent 25 days in the field sampling in as many areas as possible, and collected just over one thousand specimens. The species list for the site stands currently at 350 taxa, and continues to grow as I identify collections. Within the next year, my aim is to focus on collecting at the higher elevations, as it is challenging to get good coverage over the area. I have collected extensively within the canyons this past year. Recent work searching herbaria (RSA-POM, CAS/DS, UC/JEPS) for historical collections has produced relatively few specimens, reflecting sporadic collecting throughout the years. The majority of specimens have come from Jon Keefe, a botanist from the Los Angeles area who collected the common species in the canyons, and Verne Grant, a noted authority on the Polemoniaceae, who collected a number of Gilia species in Short Canyon. Most of the vouchers come from the three canyons within the study site, while few collections have been made at the higher elevations, as the area is relatively inaccessible and does not have any available water. The lack of available water has been a significant impediment to more extensive exploration at the higher ridges and peaks. Floras are dynamic, and floristic studies are never complete, but conducting and publishing such studies is vital to the understanding and conserving of our botanical heritage. The floristic documentation of this poorly collected area will aid in our understanding of California’s floristic province boundaries, and will also be of value in managing and conserving the region’s botanical diversity. The prospective date of completion for this project is Fall of 2004. It is my hope that this study will form a foundation upon which more extensive botanical work may be conducted in the Owens Peak area, and the Southern Sierra Nevada in general. ACKNOWLEDGMENTS I would like to thank Southern California Botanists, who so generously provided me with the Southern California Botanists Student Research Grant in 2002. With these funds I have been able to purchase a GPS, altimeter, and four Rite-in-the-Rain® field notebooks, which have aided me in CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 43 acquiring the information I need for relevant label data. 1 would like to thank my committee for their time and advice, and thanks would not be complete without acknowledging all of the generous volunteers who have helped me tremendously with fieldwork. LITERATURE CITED Hartman, R.L., and L. Constance. 1988. A new Lomatium [ shevockii ] (Apiaceae) from the Sierran crest of California. Madrono 35(2):2 1 - 1 28. Sanders, A.C., D.L. Banks, and S. Boyd. 1997. Note: rediscovery of Hemizonia mohavensis (Asteraceae) and addition of two new localities. . Madrono 44(2): 197-203. Shevock, J.R. 1988. New, rare and geographically interesting plants along the crest of the Southern Sierra Nevada, California. Pp. 161-166 in Hall, C.A. Jr., and V. Doyle- Jones (eds.) Plant Biology of Eastern California. Natural History of the White-Inyo Range Symposium Volume 2. University of California, White Mountain Research Station, Los Angeles. Shevock, J.R., B. Ertter, and J.D. Jokerst. 1989. Monardella beneolens (Lamiaceae), a new species from the crest of the Southern Sierra Nevada, California. Madrono 36(4):27 1-279. Twisselman, E.C. 1967. A flora of Kern County California. The Wasmann Journal of Biology 25(1,2). 44 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] "Challenging” terrain of the Owens Peak eastern watershed, as it is referred to by Naomi Fraga, in the previous article on the vascular flora there. The ruggedness of the Sierra is evident here along the divide between Indian Wells Canyon and Short Canyon, taken in Naomi's study area. This photograph was taken March 29, 2003. — Carl Wishner, Editor. Photographer CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 45 VASCULAR PLANT CHECKLIST FOR THE UNIVERSITY OF CALIFORNIA NATURAL RESERVE SYSTEM’S SAN JOAQUIN FRESHWATER MARSH RESERVE Peter A. Bowler Department of Ecology and Evolutionary Biology University of California, Irvine Irvine, California 92697-2525 pabowler@uci.edu Mark A. Elvin UCI Arboretum and Herbarium (IRVC) University of California, Irvine Irvine, California 92697-1459 melvin@uci.edu ABSTRACT: The University of California Natural Reserve System’s San Joaquin Freshwater Marsh Reserve was established in 1969, and is administered by the University of California, Irvine campus. The Reserve comprises 81.75 ha (202 ac) and is located in the City of Irvine, Orange County; 75 km (45 mi) southeast of Los Angeles, 30 km (20 mi) west of the Santa Ana Mountains; 2 km (1.25 mi) upstream from Upper Newport Bay, adjacent to the Irvine campus. The elevation of the site is 2-3 m (7 to 10 ft), it receives 30 cm (12 in) of precipitation per year, and average temperatures are 29° C (86° F) in September, 4° C (40° F) minimum in January, with an annual mean of 17° C (62° F). The San Diego Creek flood control channel crosses the Reserve along its southwestern boundary. Approximately 20 ha (50 ac) of the Reserve have been restored into twelve palustrine habitats dominated by bulrushes ( Schoenoplectus americanus , S. californicus and Bolboschoenus maritimus), and remainder is predominantly a cattail marsh. Nineteen acres (7.7 ha) surrounding the wetlands are a buffer zone between future development on the University’s North Campus and the Reserve. The buffer zone is currently being restored to a coastal sage scrub community, and seven vemal pools have been created at its base using inoculum from pools taken on University and adjacent Irvine Company land. Including 3 1 species that have been deliberately introduced for restoration or habitat enhancement purposes. The vascular plant flora includes one Pteridophyte and 188 species in 46 families of dicotyledones and 54 species in 9 families of monocotyledones. Of the total 242 species in the Reserve and buffer zone, 1 14 (47%) are non- native. Voucher specimens from the Reserve for most of the species are held in the UCI Herbarium (IRVC), a part of the UCI Arboretum. The non-native species Avena sativa (Poaceae), Washingtonia robusta, Phoenix dactylifera (Arecaceae), Chenopodium glaucum (Chenopodiaceae), Eucalyptus citriodora (Myrtaceae), Diospyros lycioides (Ebenaceae), Atriplex suberecta (Chenopodiaceae), Phalaris canariensis (Poaceae), and Callistemon citrinus (Myrtaceae) are new records for Orange County. KEYWORDS: University of California Natural Reserve System, San Joaquin Freshwater Marsh Reserve, Irvine, Orange County, California, Wetland Restoration, Coastal Sage Scrub Restoration, Vemal Pool Creation, California Coastal Conservancy 46 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] INTRODUCTION The University of California Natural Reserve System’s San Joaquin Freshwater Marsh Reserve is located in the City of Irvine (Orange County) adjacent to the University of California (UCI) campus, located approximately 2 km from Newport Back Bay. The Reserve comprises 81.75 ha (202 ac) and includes the channelized San Diego Creek that passes through it. Pollen in cores obtained from the Reserve indicate that several times in the last 6,000 years, the site was a salt marsh, switching back and forth between freshwater and salt marsh plant species as the ocean level rose and fell (Davis, 1992; Davis, Jirikowic, and Kalin, 1992). The Marsh is a remnant wetland on a filled canyon of the Santa Ana River, inland from the Newport Bay Gap; one of five such canyons marking the shifting course of the Santa Ana River in the past. An historic dam on San Diego Creek, an artificial stream draining urban and agricultural runoff from Irvine and surrounding areas, caused several meters of sediment to collect in the southern portion of the Reserve. The dam was constructed about 1937 to protect salt desiccation ponds in Newport Back Bay, and was dismantled in the late 1950s. The sediment behind the dam raised the ground level above natural wetland elevations, and the site was actually farmed as upland for several decades. The bluffs surrounding the wetlands were similarly used for agriculture, and the Reserve experienced cattle grazing until the University purchased it in 1969. The California Coastal Conservancy sponsored restoration efforts in 1998 on 18.6 ha (46 ac) of the Reserve, allowing the creation of 1 1 palustrine habitats at historic pre-dam elevations. These ponds have about 40 percent of their area designed to provide permanent deep water habitat, with the remainder being shallow or seasonally dry shelves dominated by Bolboschoenus maritimus, Schoenoplectus americanus and S. californicus. An additional 2.4 ha (6 ac) are a created marsh as part of a mitigation project. The Conservancy has also supported creating coastal sage scrub as a complementary upland community along 7.7 ha (19 ac) of bluffs embracing the wetland as part of a 50 m buffer zone between existing and future development on the University’s North Campus. The remaining 60.7 ha (150 ac) are primarily a 7yp/w-dominated seasonal wetland, with a natural soil sequence underlying it. In addition to precipitation (30 cm/year), water can be introduced to the Reserve from San Diego Creek, a well, or allowed to flow under adjacent Campus Drive from upstream sources. The Reserve System manages the site following the area’s natural hydrologic cycle so that much of the wetlands are seasonally wet or dry as the natural climate dictates. Prior to disturbance, the site was a ground water depression wetland supported by a shallow aquifer. This part of Orange County has had vast losses of wetlands including over 20,000 ha in the historic Cienega de las Ranas that once covered the Tustin Plain, the construction of the Irvine Ranch Water District’s Michelson Sewage Treatment Plant and associated facilities, the placement of Campus Drive through the wetlands, and the channelization and construction of the San Diego Creek channel for flood control purposes. Thus the UCNRS San Joaquin Freshwater Marsh Reserve is a significant wetland remnant near the larger salt marsh wetlands in the State’s Upper Newport Bay Ecological Reserve. The UCNRS uses the wetlands Reserve for teaching and research, and because of controlled human access it is a true refuge, with 263 bird species and abundant other wildlife having been recorded there. The present Checklist includes vascular plant species within the Reserve and within the 50 m protected buffer zone surrounding it on the western side along University lands. The current Vascular Plant Checklist for the UCNRS San Joaquin Marsh Reserve is based upon Bowler and Wolf (1993), with many additions since the original Checklist was published a decade ago. Checklists are works in progress, and subject to change as taxa invade, are discovered, are purposely introduced, or disappear from a site. Taxa introduced deliberately as part of coastal sage scrub restoration in the buffer zone between the Marsh Reserve and North (UCI) Campus development include Chlorogalum pomeridianum (Bowler, 1999), Dudleya lanceolata, D. multicaulis, D. pulverulenta, Eriogonum cinereum, Eriogonum elongation, Eriogonum CROSSOSOMA 29(2), Fall- Winter 2003 [issued August 2004] 47 fasciculatum, Ceanothus megacarpus, Rhus integrifolia, Leymus condensatus, Malosma laurina, Mimulus aurantiacus, Mirabilis laevis, Cylindropuntia prolifera, Opuntia litloralis, Heteromeles arbutifolia, Salvia mellifera, Sisyrinchium bellum, and Isomeris arborea. Rosa californica obtained from Laguna Canyon was introduced at the gate between the Reserve and UCI Arboretum, and it is now established in the buffer zone at that location, as well as having been introduced elsewhere in the Reserve. Platanus racemosa, Populus fremontii, and Quercus agrifolia have been planted both at the edge of the Marsh and in the buffer zone. In 1998, two vernal pools were created along the edge of the marsh, and these were inoculated with mud from pools on the main UCI campus that included, among others, the following taxa new to the marsh flora: Psilocarphus brevissimus, Plagiobothrys acanthicarpus, Juncus bufonius, Lythrum hyssopifolium, Lythrum Iribracteatum, Plantago elongata, Veronica peregrina ssp. xalapensis, and Eleocharis macrostachya. Five additional pools were established during November 2002, using the same inoculum. In both cases, the source material was from the University of California, Irvine main campus, or other local sites. In addition to the vernal pool flora, the Riverside fairy shrimp, Branchinecta lindahlii (Packard), was successfully established in them. Adrian Wolf found Atriplex coulteri , the rare native Coulter’s saltbush, on the UCI Landfill that abuts the Reserve. It is included in the Checklist because it may have been found within the buffer zone that includes part of the landfill. This rare species is on the California Native Plant Society’s IB list, “species that are rare throughout their range and occur primarily within California (Roberts 1998). Centromadia parryi ssp. australis, southern tarplant, reported from the Marsh in the initial checklist, is a special concern taxon tracked by the California Natural Diversity Data Base. Juncus balticus, the native wire rush, Ludwigia peploides, yellow waterweed, Cyperus odoratus, fragrant umbrella-sedge, Pentagramma triangularis, silverback fern, Lemna minuta (overlooked but in IRVC), Chaemaesyce albomarginata (also overlooked but in IRVC), Asclepias fascicularis, the narrow- leaf milkweed, Pectocarya linearis ssp.ferocula, slender Pectocarya, Cardamine oligosperma, few- seeded bittercress, Opuntia X occidentals, western prickly pear, Daucus pusillus, rattlesnake weed. Datura stramonium, Jimson weed, Cuscuta californica, Cuscuta subinclusa, and Lepidium lasiocarpum were also added to the list. Since 1993, the European garland or crown daisy, Chrysanthemum coronarium, has established a large stand along San Diego Creek in the Reserve, and has some presence on the adjacent dikes in the Marsh. Other exotics added to the list are Urtica urens, the Old World dwarf nettle, Oxalis pes-caprae, Bermuda buttercup, Chenopodium glaucum, glaucous leaved saltbush, Erodium botrys, long-beaked filaree, Marrubium vulgare, common horehound, Cotula australis, Australian brass-buttons, Melilotus officinalis, yellow sweet- clover, Malva sylvestris, high mallow, Callistemon citrinus, bottlebrush, Schinus terebinthifolius, Brazilian pepper tree, Gnaphalium luteo-album, weedy cudweed, Beilis perennis, English daisy, Avena sativa, cultivated oat, Lobularia maritima, sweet-alyssum, Washingtonia robusta, Mexican fan palm, Phoenix dactylifera, date palm, Eucalyptus citriodora, Callistemon citrinus, lemon bottle brush, Diospyros lycioides, Atriplex suberecta, halberd-leaved saltbush, Phalaris canariensis, canary grass, Lamarckia aurea, goldentop, and Lycospersicon esculentum, the South American tomato. See Bowler and Wolf (1994) for a discussion of invasive plants in the Reserve. In a September, 2002 survey of the Irvine Ranch Water District (IRWD) mitigation areas across Campus Drive from the UCNRS San Joaquin Marsh Reserve, Tara Schoenwetter recorded a number of taxa not yet occurring the Reserve. Because of their establishment in a site adjacent the Reserve, I have listed them as likely invaders in the Checklist with the location identied as IRWD. The IRWD taxa include the natives Chenopodium berlandieri, pitseed goosefoot, Salix laevigata, red willow, and Paspalum distichum, knot grass, and the exotics are Lepidium draba, the Eurasian heart-podded hoary-cress, Chamaesyce serpens, the South American annual rattlesnake spurge, Chamaesyce maculata, spotted spurge from the eastern U.S., Oenothera speciosa, showy-evening primrose from the central US and Mexico, Epilobium sp., Rumex conglomeratus, whorled dock from Europe, Polygonum persicaria, lady’s-thumb from Europe, Plantago major, common 48 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] plantain from Europe, Pennisetum clandestinum, Kikuyu grass, and Lepidium latifolium, the Eurasian broad-leaved peppergrass that has recently invaded the Irvine Ranch Water District’s property across Campus Drive. Lepidium latifolium is the primary species of concern for the Reserve, since the other exotics are already ubiquitous in Orange County. It is hoped that exotics such as Acacia, Eucalyptus, Myoporum, Phoenix, Washingtonia , Celtis, Schinus, Callistemon, and Disospyros will be eliminated from the Reserve, and an on-going aggressive eradication effort is being conducted upon tamarisk, fennel, poison hemlock, artichoke, star thistle, castor bean, giant reed, and black mustard. Taxa will continue to be added to the Checklist as they are discovered on the Reserve. Including 3 1 species deliberately introduced for restoration or habitat enhancement purposes, the vascular plant flora consists of one Pteridophyte and 188 species in 46 families of dicotyledones, and 54 species in 9 families of monocotyledones. Of the total 242 species in the Reserve and buffer zone, 1 14 (47%) are non-native. Nine non-native and three native species apparently not in Reserve occur on adjacent IRWD property. The numbers of native and non-native taxa in the ten largest families are presented in Table 1, the genera and species numbers in the Reserve with their counterparts in the Orange County flora appear in Table 2, and the number of native and non-native species in each family occurring on the Reserve comprise Table 3. CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 49 Table 1. The number of species in the ten largest families in Orange County (Roberts 1998) compared with those same families as represented in the UC Natural Reserve System’s San Joaquin Marsh Reserve. The last column entry for a family indicates the current flora including deliberate restoration introductions. Family Orange County species (native; non-native) San Joaquin Marsh Reserve species (native; non-native) including restoration introductions Asteraceae 186 (121; 65) 53 (32; 21) 54 (33; 21) Poaceae 124 (50; 74) 30 (4; 26) 31 (5; 25) Fabaceae 84 (49; 35) 6 (1; 5) Brassicaceae 47 (22; 25) 13 (4; 9) Chenopodiaceae 41 (26; 15) 16 (7; 9 Cyperaceae 34 (31; 3) 8 (8; 0) 10(10; 0) Scrophulariaceae 38 (29; 9) 1 (0; 1) 3(2; 1) Polygonaceae 32 (25; 7) 4 (2; 2) 7 (5; 2) Apiaceae 30 (22; 8) 5 (2; 3) Onagraceae 28 (24; 4) l ( 1 ; 0) Table 2. A summary of the Angiosperm flora of Orange County (OC) (adapted from Roberts 1998) and the UCNRS San Joaquin Marsh Reserve. The Marsh figures in brackets represent the introduced restoration taxa added into the broader flora. Genera Species (all) native non- native OC Marsh OC Marsh OC Marsh OC Marsh Angios. 528 1 44[ 1 65] 1,156 204[236j 773 97[ 1 28] 383 1 1 2[ 1 14] Dicots 426 1 1 2[ 1 30] 927 1 62[ 1 88] 634 79[ 103] 293 83[85] Monocots 102 35[36] 229 49[54] 139 20[25] 90 29 Table 3. The families of Angiosperms and their native and non-native representatives in the UCNRS San Joaquin Marsh Reserve. The first three columns following the family are the flora not reflecting deliberate restoration (RI) or vernal pool creation (VPI) introductions. The final column on the right indicates the current flora, including them. Flora Totals (without introductions) species (native; non-native) Introductions Flora Totals (with introductions) species (native; non-native) GROUP Family DICOTS Adoxaceae Aizoaceae 1 (1;0) 4(1; 3) Amaranthaceae 3 (1; 2) Anacardiaceae 3 (1; 2) RI 2 native 5 (3; 2) Apiaceae 5 (2; 3) Asclepiadaceae l (1; 0) Asteraceae 53 (32; 21) VPI 1 native 54 (33; 21) Boraginaceae 3 (3; 0) VPI 1 native 4 (4; 0) Brassicaceae 13 (4; 9) 50 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Cactaceae id; 0) RI 2 native 3 (3; 0) Capperaceae 0 (0; 0) RI 1 native 1(1,0) Caryophyllaceae 3(1:2) Chenopodiaceae 16 (7; 9) Convolvulaceae 3(2; 1) Crassulaceae 1 ( 1; 0) RI 3 native 4 (4; 0) Cucurbitaceae 1 ( 1 ; 0) Cuscutaceae 3 (3; 0) Ebenaceae 1(0:1) Euphorbiaceae 5 (3; 2) Fabaceae 6(l;5) Fagaceae 0 (0; 0) RI 1 native 1(1:0) Frankeniaceae 1 0:0) Geraniaceae 3 (0; 3) Lamiaceae 2 (0; 2) RI 1 native 3(1; 2) Linaceae 1 (0; 1) Lythraceae 0 (0; 0) VPI 2 non-native 2 (0; 2) Malvaceae 4(1:3) Myoporaceae 1 (0; 1) Myrtaceae 2 (0; 2) Nyctaginaceae 0 (0; 0) RI 1 native 1(1:0) Onagraceae 1(1:0) Oxalidaceae 1(0; 1) Plantaginaceae 1 (1:0) VPI 1 native 2 (2; 0) Platanaceae 0 (0; 0) RI 1 native 1(1:0) Polygonaceae 4 (2; 2) RI 3 native 7 (5; 2) Portulacaceae 1(0; 1) Primulaceae 1(0; 1) Rhamnaceae 0 (0; 0) RI 1 native 10:0) Rosaceae 0 (0; 0) RI 2 native 2 (2; 0) Rubiaceae 1(0; 1) Salicaceae 3 (3; 0) RI 1 native 4 (4; 0) Saururaceae TTTTo) Scrophulariaceae 1(0; 1) VPI/RI 2 native 3(2; 1) Solanaceae 8 (5; 3) Tamaricaceae 1(0; 1) Urticaceae 2(1; 1) MONOCOTS Arecaceae 3 (0; 3) Cyperaceae 8 (8; 0) VPI 1 native 9 (9; 0) Iridacaceae 0 (0; 0) RI 1 native l ( l ; 0) Juncaceae 1 0:0) VPI 1 native 2 (2; 0) Lemnaceae 1(1:0) Liliaceae 3(2; 1) RI 1 native 4(3; 1) Poaceae 30 (4; 26) RI 1 native 31 (5; 25) Potamogetonaceae 1(1:0) Typhaceae 3(3;0) CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] ACKNOWLEDGMENTS 51 Adrian Wolf and Tara Schoenwetter provided determinations or other information about some of the new records. Andy Sanders assisted in several determinations. I thank my Independent Study students, particularly Jenny Liou, for assistance in collecting vouchers of several dozen species, and for keeping the restoration areas alive until establishment. This study was supported in part by Grants 01-130 and 99-039 from the California Coastal Conservancy. I am especially grateful to my wife, Christine Hager, and her mother LaVeme Hager for help with data entry and analysis. This work was performed at the University of California Natural Reserve System San Joaquin Freshwater Marsh Reserve, and technical assistance was provided by the University of California, Irvine Office of Natural Reserves. In 1981, staff of the former UCI Museum of Systematic Biology, primarily Fred Roberts, Jr., Gordon Marsh and Ann McGee collected vouchers of many taxa. These, and other accessioned herbarium collections indicated in the Checklist reside in the University of California, Irvine Herbarium (IRVC), a part of the UCI Arboretum (contact pabowler@uci.edu for access to the Herbarium). Most of the taxa on the Checklist (178 species, or 75 percent) are represented by vouchers in IRVC, with the eventual goal of vouchering at least 90 percent of the flora. A continuously updated Checklist as modified from the original 1993 Crossosoma publication may be accessed online at UCNRS San Joaquin Marsh Reserve website http://nrs.ucop.edu/reserves/sjfm.html. The lichen flora of the Reserve (Bowler and Riefner 1990) also appears on Reserve webpage. LITERATURE CITED Bowler, P.A. 1999. Transplantation of Understory Bulbs, Grasses, Lichens and Bryophytes in Coastal Sage Scrub Restoration. Ecological Restoration 17:82-83. Bowler, P.A., and A. Wolf. 1993. Vascular Plants of the San Joaquin Freshwater Marsh Reserve. Crossosoma 19(1 ):9-30. Bowler, P.A., and R. Riefner, Jr. 1990. A Preliminary Lichen Checklist for the University of California, Irvine, Campus and the San Joaquin Wetlands. Crossosoma 16(6): 1-10. Bowler, P.A. , and A. Wolf. 1994. Exotic Plants in Mediterranean Climate Wetlands. Orange County, California: A Case Study. Crossosoma 20(1 ):75-84. Bowler, P.A., A. Wolf, and L. Bradley. 1995. A Checklist of the Wetland Indicator Species in Orange County, California. Crossosoma 21(1): 1-39. Davis, O.K. 1992. Rapid Climatic Change in Coastal Southern California Inferred from Pollen Analysis of San Joaquin Marsh. Quaternary Research 37:89-100. Davis, O.K., J. Jirikowic, and R.M. Kalin. 1992. Radiocarbon Record of Solar Variability and Holocene Climatic Change in Coastal Southern California. 14 pp. in Redmond, K.T. (ed.). 1992. Proceedings of the Eigth Annual Pacific Climate (PACLIM) Workshop, March 10-13. 1991 : California Department of Water Resources, Interagency Ecological Studies Program Technical report 31. Hickman, J.C. (ed.). 1993. The Jepson Manual: Higher Plants of California. The University of California Press. Berkeley, California. Munz, P.A. 1968. A California Flora with Supplement. University of California Press, Berkeley, California. Reed, P.B., Jr. 1988a. National list of plant species that occur in wetlands: national summary. U.S. Fish & Wildlife Service, Biol. Rep. 88(24). Reed, P.B., Jr. 1988b. National list of plant species that occur in wetlands: California (Region 0). U.S. Fish and Wildl. Serv. Biol. Rep. 88(26.10). Roberts, F.M., Jr. 1998. A Checklist of the Vascular Plants of Orange County, California. Second Edition. F.M. Roberts Publications, Encinitas, California. 52 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] The Vascular Plant Checklist for the University of California Natural Reserve System’s San Joaquin Freshwater Marsh Reserve PTERIDOPHYTA — Ferns and Fern-Allies and ANTHOPHYTA — Flowering Plants The following symbols are used to indicate in which habitat most species might be anticipated to occur W - wet, standing water; M = moist, saturated soils; I = intermediate; D = dry, water may be limiting. There are undoubtedly overlaps in areas of occurrence. WI = National Wetlands Indicator Species (Reed 1988a, 1988b; Bowler, Wolf and Bradley 1995). The symbol + indicates that a voucher collection of the species from the UCNRS San Joaquin Freshwater Marsh Reserve is in the UCI Herbarium (IRVC) curated by the UCI Arboretum. RI appearing before a scientific name = a deliberate introduction for restoration or habitat enhancement purposes. VPI appearing before a scientific name indicates a deliberate introduction in created vernal pools adjacent the Marsh. IRWD appearing before a scientific name = a species not yet recorded from the Reserve that occurs on adjacent IRWD property . Flowering times are primarily taken from Munz (1968) and scientific nomenclature follows the Jepson Manual (Hickman 1 993), except where superseded names are given in the Jepson Online Interchange [updated by the Editor]. PTERIDOPHYTA — FERNS AND FERN-ALLIES Pteridaceae — Brake Family Pentagramma triangularis (Kaulfuss) Yatskievych, Windham & Wollenweber Silverback Fem D ANTHOPHYTA — FLOWERING PLANTS DICOTYLEDONES — DICOTS Adoxaceae — Honeysuckle Family Sambucus mexicana C. Presl +, WI Blue Elderberry Spring-Fall I, D Aizoaceae — Fig-Marigold or Carpet- Weed Family *Malephora crocea (Jacq.) Schwantes Croceum Ice Plant S Africa Summer D *Mesembryanthemum crystallinum L. + Crystalline Ice Plant S Africa Mar-Oct D CROSSOSOMA 29(2), Fall- Winter 2003 [issued August 2004] 53 *Mesembryathemum nodijlorum L. + Slender-leaved Ice Plant S Africa Apr-Nov D Sesuvium verrucosum Raf. +, WI Western Sea-Purslane Apr-Nov D Amaranthaceae — Amaranth Family *Amaranthus albus L. +, WI Tumbleweed Trop. Am. May-Oct D Amaranthus blitoides S. Watson +, WI Prostrate Pigweed Europe May-Nov D * Amaranthus retroflexus L. WI Rough Pigweed Trop. Am. Jun-Nov I, D Anacardiaceae — Cashew or Sumac Family Rhus integrifolia (Nuttall) Brewer & S. Watson +, RI Lemonade berry Malosma laurina (Nuttall) Abrams +, RI Feb-May I, D Laurel sumac Jun-Jul I, D *Schinus terebinthifolius Raddi + Brazilian peppertree I Toxicodendron diversilobum (Torrey & A. Gray) E. Greene + Western Poison-Oak Apr- May LD Apiaceae (Umbelliferae) — Carrot Family *Apium graveolens L. +, WI Celery Apiastrum angustifolium Nuttall Eurasia May-Jul M, I Mock Parsley Mar-Apr D *Conium maculatum L. +, WI Poison-Hemlock Europe Apr-Sep M, I Daucus pusillus Michaux + Rattlesnake Weed Feb-Apr D *Foeniculum vulgare Miller +, WI Fennel Europe May-Sep M, I Asclepiadaceae — Milkweed Family Asclepias fascicularis Decne. WI Narrow-leaf Milkweed Apr-Jun I,D Asteraceae (Compositae) - - Sunflower Family *Acroptilon repens (L.) DC. + Russian Knapweed Europe May-Sep M Ambrosia acanthicarpa Hook. + Annual Bur-sage Ambrosia psilostachya DC. +, WI Aug-Nov M, I, D Western Ragweed Jul-Nov I,D 54 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] *Anthemis cotula L. +, WI Dog-fennel, Mayweed Europe Apr-Sep M, I, Artemisia biennis Willd. +, WI Biennial Sagewort Aug-Oct I Artemisia califomica Lessing + Coastal Sagebrush Aug-Dec D Artemisia douglasiana Besser +, WI Mugwort Jun-Oct I Artemisia dracunculus L. + Dragon Sagewort, Tarragon Aug-Oct I, D Symphyotrichum divaricatum (Nutt.) G.L. Nesom [<= Aster subulatus Michaux var. ligulatus Stunners] +, WI Slender Aster Jul-Oct I, D Baccharis emoryi A. Gray. +, WI Emory’s Baccharis Aug-Dec M, I Baccharis pilularis DC. + Coyote Bush, Chaparral Broom Aug-Dec M, I Baccharis salicifolia (Ruiz Lopez & Pavon) Pers. +, WI Mule Fat Mar-Jun W, N *Bellis perennis L. + English Daisy Europe Early spring W,f *Centaurea melitensis L. + Tocalote Europe May-Jun M, I, Centromadia parryi E. Greene ssp. australis (Keck) 1 Baldwin [<= Hemizonia parryi E. Greene ssp. australis Keck] 1 + Southern Tarplant June-Sep I, D Chamomilla suaveolens (Pursh.) Rydb. +, WI Common Pineapple Weed May-Aug I, D * Chrysanthemum coronarium Linn. + Garland or Crown Daisy Europe May-Jul I *Cirsium vulgare (Savi) Ten. +, WI Bull Thistle Europe Jun-Sep I *Conyza bonariensis (L.) Cronq. + Flax-Leaved Horseweed S Am. Jun-Sep D Conyza canadensis (L.) Cronq. Horseweed Jun-Sep I, D Conyza coulteri A. Gray var. virgata (Benth.) Gray WI Coulter's Horseweed Jun-Sept M, I, Corethrogyne filaginifolia (Hooker & Amott) Nutt. [< c= Lessingia f. M. A. Lane var./.] Virgate Cudweed Aster Jul-Oct I, D *Cotula australis (Sieber) Hook. f. + Australian Brass-Buttons Australia Mar-May D *Cotula coronopifolia L. +, WI African Brass-Buttons S Africa Feb-Jul M, I *Cynara cardunculus L. + Cardoon, Globe Artichoke Medit. May-Jul I, D Eclipta prostrata (L.) L. False Daisy All mo. M Ericameria palmeri (A. Gray) H.M. Hall + Goldenbush Aug-Dec D CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 55 Euthamia occidentalis Nuttall +, Wl Western Goldenrod Jul-Nov M Filago californica Nutt. + California Filago, Fluffweed Mar-Jun I Gnaphalium californiacum DC. + California Everlasting Jan-Jul I, D Gnaphalium canescens DC. ssp. microcephalum (Nuttall) Stebbins & Keil + White Everlasting Jul-Oct I *Gnaphalium luteo-album L. + Weedy Cudweed Eurasia Apr-Jul D Gnaphalium palustre Nutt. WI Lowland Cudweed May-Oct M Grindelia camporum E. Greene var. bracteosum (J. Howell) M.A. Lane +, WI Big Gumplant Mar-Sep M, I, D Helianthus annus L. +, WI Western Sunflower Feb-Oct M, I Hemizonia fasciculata (DC.) Torr. & A. Gray + Fascicled Tarweed May-Sep I, D Heterotheca grandiflora Nutt. + Telegraph Weed Jan-Dec I, D Heterotheca sessiliflora (Nutt.) Shinn, ssp. echioides (Benth.) Semple Goldenaster Jul-Nov I, D *Hypochaeris glabra L. + Smooth Cat’s-Ear Europe Apr-Jun I, D Isocoma menziesii (Hook. & Am.) G. Nesom var. vernonioides (Nutt.) G. Nesom +, WI Coastal Goldenbush Apr-Dee D *Lactuca serriola L. +, WI Prickly or Wild Lettuce Europe May-Sep M, I, D *Picris echioides L. +, WI Bristly Ox-Tongue Europe Jun-Dec M, I Pluchea odorata (L.) Cass. +, WI Salt Marsh-Fleabane Jul-Nov M, I Psilocarphus brevissimus Nutt. var. brevissimus +, VPI, WI Dwarf Woolly Heads Apr-May VP *Pulicaria paludosa Link + Spanish Sunflower Europe Aug-Sep M *Senecio vulgaris L. +, WI Common Groundsel Eurasia Spring-Summer I, D *Silybum marianum (L.) Gaertner + Milk Thistle Medit. May-Jul I *Soliva sessilis Ruiz Lopez & Pavon Coast Soliva S Am. Apr-Jul M *Sonchus asper (L.) Hill. +, WI Prickly Sow Thistle Europe Most mos. M, I *Sonchus oleraceus L. +, WI Common Sow Thistle Europe Most mos. M, I *Sonchus tenerrimus L. WI Slender Sow Thistle Apr-Jul 9 56 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Stebbinsoseris heterocarpa (Nutt.) Chambers Derived Stebbinsoseris + Stephanomeria virgata Benth. Tall Wreath-Plant Xanthium strumarium L. +, WI Cocklebur Apr-Map Jul-Oct Jun-Oct Boraginaceae — Borage Family Amsinckia menziesii (Lehm.) A. Nelson & J.F. Macbr. var. intermedia (Fischer & C. Meyer) Ganders + Rancher's Fireweed Feb-May Heliotropium curassavicum L. +, WI Salt or Alkali Heliotrope Mar-Oct Plagiobothrys acanthicarpus (Piper) I. M. Johnston WI, VPI Adobe Popcorn Flower Mar Pectocarya linearis (R. & P.) DC. ssp .ferocula (Jtn.) Thome + Slender Pectocarya Feb- Apr Brassicaceae (Cruciferae) — Mustard Family M, I, D I, D M I, D I, D I, M D *Brassica nigra (L.) Koch + Black Mustard Europe Apr-Jul *Brassica rapa L. + Field Mustard Europe May-Oct Cardamine oligosperma Torrey & A. Gray +, WI Few-Seeded Bittergrass Mar-Jul *Hirschfeldia incana (L.) Lagr.-Fossat + Shortpod/Summer Mustard Medit. Basin May-Oct *Lepidium didymum L. [<= Coronopus didymus (L.) Sm.] Lesser wart-cress Eurasia Feb- Apr *Lepidium draba L. ssp. draba [<= Cardaria draba (L.) Desv.] IRWD Heart-podded Hoary Cress Eurasia *Lepdium latifolium L. IRWD Broad-leaved Peppergrass Eurasia Lepidium lasiocarpum Torrey & A. Gray + Sand Peppergrass Lepidium nitidum Torrey & A. Gray + Mar-May May-Sep Feb-May I I D I + D D, M, I I, M D Shining Peppergrass Feb-May I Lepidium virginicum L. var. pubescens (E. Greene) Thell. +, WI Wild Peppergrass *Lobularia maritima (L.) Desv. + Mar-Aug I Sweet-Alyssum Medit. Spring-Fall I, D *Raphanus sativus L. + Wild Radish Medit. Feb-Jul I * Sisymbrium irio L. + London Rocket Europe Feb-May M, I, D CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 57 Cactaceae — Cactus Family Cylindropuntia prolifera (Engelm.) F.M. Knuth [<= Opuntia p. Engelm.J +, RI Coastal Cholla Late spring D Opuntia littoralis (Engelm.) Cockerell +, RI Coastal Prickly Pear May-Jun I, D Opuntia X occidentalis Engelm. + Western Prickly-Pear May-Jul D Capparaceae — Caper Family Isomeris arborea Nutt. +, RI Bladderpod All year D Caryophyllaceae — Pink Family Spergularia marina (L.) Griseb. +, WI Salt-marsh Sand Spurry Mar-Sep I, D *Spergularia villosa (Pers.) Cambess. Villous Sand Spurry S Am. Apr-Jul I, D *Spergularia cf. rubra (L.) J.S. Presl & C. Presl + Red Sand Spurry Europe Summer I, D Chenopodiaceae - Goosefoot Family Atriple x canescens (Pursh) Nutt. +, WI Fourwing Saltbush; Shadscale May-Jul D Atriplex coulteri (Moq.) D. Dietr. WI Coulter’s Saltbush Mar-Oct D Atriplex lentiformis (Torrey) S. Watson ssp. lentiformis +, WI Big Saltbrush May-Aug S * Atriplex rosea L. +, WI Tumbling Oracle Eurasia Jul-Oct I, D * Atriplex semibaccata R.Br. +, WI Australian Saltbush Australia Apr-Dee M, I, D * Atriplex suberecta I. Verd. + Halberd-leaved Saltbrush Australia Mar-Jul M Atriplex triangularis Willd. +, WI Halberd-leaved Saltbrush Jun-Nov M *Bassia hyssopifolia (Pallas) Kuntze +, WI Five-hook Bassia Eurasia Jul-Oct I, D *Beta vulgaris L. +, WI Beet Europe Jul-Oct M, I *Chenopodium album L. +, WI Lamb's Quarters; Pigweed Europe Spring-Summer M, I *Chenopodium ambrosioides L. +, WI Mexican Tea Trop. Am. Jun-Oct I Chenopodium berlandieri Moq. IRWD Pitseed Goosefoot Mar-Sept I, D 58 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004) *Chenopodium glaucum L. + Eurasia *Chenopodium murale L. +, Nettle-leaved Goosefoot Europe Salicomia virginica L. +, Common Woody Pickleweed *Salsola tragus L. + Russian Thistle Eurasia Sueda esteroa W. Ferren & S. Whitmore Estuary Sea-Blite Spring? I WI Mar-Jul I WI Aug-Nov M Jul-Oct I, D Jul-Oct I Convolvulaceae — Morning-Glory Family Calystegia sepium (L.) R.Br. +, WI Hedge Bindweed Spr-Sum D *Convolvulus arvensis L. + Orchard Morning-Glory Europe May-Oct I, D Cressa truxillensis Kunth +, WI Alkali Weed May-Oct I Crassulaceae ■ — Stonecrop Family Crassula connata (Ruiz Lopez & Pavon) A. Berger +, WI Pygmy Stonecrop Feb-May I, D Dudleya lanceolata (Nutt.) Britton & Rose +, RI Lance- leaf Dudleya Spring D Dudleya multicaulis (Rose) Moran +, RI Many-Stemmed Dudleya Late Spring D Dudleya pulverulenta (Nutt.) Britton & Rose +, RI Chalky Live-forever Early Summer D Cucurbitaceae — Gourd Family Cucurbita foetidissima Kunth. + Calabazilla; Stinking Gourd Jun-Oct I, D Cuscutaceae — Dodder Family Cuscuta salina Engelm. Pickleweed Dodder Jul-Nov M, I Cuscuta californica Durand & Hilg. Dodder May-Jul I, D Cuscuta subinclusa Durand & Hilg. + Dodder May-Jul W, M, I Ebenaceae — Ebony Family *Diospyros lycioides Desf. + S. Africa Mar-Aug I, D CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 59 Euphorbiaceae — Spurge Family Chamaesyce albomarginatus (Torrey & A. Gray) Small Rattlesnake Spurge *Chamaesyce maculata IRWD Spotted Spurge E USA *Chamaesyce prostrata (Aiton) Small Prostrate Euphorbia S Am. *Chamaeysce serpens (Kunth) Small IRWD Annual Rattlesnake Spurge S Am. Chamaesyce serpyllifolia (Pers.) Small var. hirtula (S. Watson) Koutnik Thyme-Leaved Spurge Aug-Oct Croton setigerus Muell. Arg.[<= Eremocarpus s. (Hook.) Benth.] Spring-Summer Spring-Summer Aug-Sep Spr-Summer Dove Weed May-Oct I, D *Ricinus communis L. +, WI Castor-bean Europe Most mos. I Fabaceae (Leguminosae) — Legume Family * Acacia longifolia (Andrews) Willd. + Sydney Golden; Golden Wattle Australia Spring D Lotus scoparius (Nutt.) Ottley + Coastal Deerweed; CA Broom *Medicago polymorpha L. + Mar-Aug I California Burclover Medit. Mar-Jun I *Melilotus albus Medikus +, WI White Sweetclover Eurasia May-Sep M, I *Melilotus indica (L.) All. +, WI Yellow Sweetclover Medit. Apr-Oct M, I *Melilotus officinalis (L.) Pall. + Sweetclover Fagaceae — Oak Family Quercus agrifolia Nee var. agrifolia +, RI Coast Live Oak; Encina Spring I, D Frankemaceae — Frankenia Family Frankenia salina (Molina) I.M. Johnston +, Alkali Heath Geraniaceae *Erodium botrys (Cav.) Bertol. + Long-Beaked Filaree *Erodium cicutarium (L.) L'Her. + Red-Stemmed Filaree Eurasia *Er odium moschatum (L.) L'Her. + White-Stemmed Filaree Europe WI Jun-Oct M - Geranium Family Feb-May D Feb-May I, D Feb-May I + D D D D + 60 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Lamiaceae (Labiatae) — Mint Family *Lamium amplexicaule L. Dead Nettle Eurasia Apr-Sep D *Marrubium vulgare L. + , WI Common Horehound Europe Spr-Sum I Salvia mellifera E. Greene +, RI Black Sage Spring D Linaceae — Flax Family *Linum usitatissimum L. + Common Flax Europe Apr-May D Lythraceae — Loosestrife Family *Lythrum hyssopifolium L. WI, VPI Hyssop Loosestrife Europe (Spring-early summer) VP *Lythrum tribracteatum Sprengel Three-bracted Loosestrife Europe (Spring-early summer) VP Malvaceae ■ — Mallow Family *Abutilon theophrasti Medikus + , WI Velvet Leaf S Asia Jul-Aug 7 *Malva parviflora L. + Cheeseweed Eurasia Most mos. M *Malva sylvestris L. + High mallow Europe Spring M, I, D Malvella leprosa (Ortega) Krapov. + Alkali-Mallow; Whiteweed May-Oct M, I Myoporaceae - - Myoporum Family *Myoporum laetum Forster f. + Myoporum NZ Spring M Myrtaceae — Myrtle Family *Callistemon citrinus (Curtis) Skeels + Lemon Bottlebrush Australia Summer D * Eucalyptus citriodora Hook. + Australia Spring I, D Nyctaginaceae — Four O’Clock Family Mirahilis laevis (Benth.) Curran var. crassifolia (Choisy) Spellenb. [<= M. californica A. Gray; M. c. var. c.] +, RI California Wishbone Bush Spring D CROSSOSOMA 29(2), Fall- Winter 2003 [issued August 2004] 61 Onagraceae — Evening Primrose Family Epilobium sp. IRWD, WI Summer-Fall I, D Ludwigia peploides (Kunth) Raven +,WI Yellow Waterweed Spr-Summer W *Oenothera speciosa Nutt. IRWD Showy-white Evening Primrose c US, Mex. ? D Oxalidaceae — Oxalis Family *Oxalis pes-caprae L. + Bermuda Buttercup S Africa Late fall-early summer D Plantaginaceae — Plantain Family Plantago elongata Pursh VPI, WI California alkali plantain Late Spring I, D Plantago erecta E. Morris + California plantain Spring D *Plantago major L. IRWD, WI Common plantain Europe Late Spring-Summer? M, I, Platanaceae — Sycamore or Plane Tree Family Platanus racemosa Nutt. +, RI, WI Western Sycamore Spring M, I Polygonaceae — Buckwheat Family Eriogonum fasciculatum Benth. +, RI California Buckwheat Late Spring-Early Fall D Eriogonum cinereum Benth. +, RI Gray Coast Buckwheat Late Spring D Eriogonum elongatum Benth. +, RI Long-stemmed Buchwheat Late Spring D *Polygonum arenastrum Boreau +, WI Common Knotweed, Doorweed Eurasia May-Nov I Polygonum amphibium L. var. emersum Michaux Water Smartweed; Kelp Jul-Sep W, N Polygonum lapathifolium L. WI Willow Smartweed Jun-Oct M *Polygonum persicaria L. IRWD, WI Lady’s Thumb Europe ? M *Rumex conglomeratus Murr. IRWD, WI Whorled Dock Europe Spring-Summer M *Rumex crispus L. +, WI Curly Dock Eurasia Mar-Jun I 62 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Portulacaceae — Purslane Family *Portulaca oleracea L. WI Common Purslane Europe May-Sep S Primulaceae — Primrose Family *Anagallis arvensis L. +, WI Scarlet Pimpernel Europe Mar-Jul I Rhamnaceae — Buckthorn Family Ceanothus megacarpus Nutt. ssp. megacarpus +, RI Bigpod Lilac Spring D Rosaceae — Rose Family Heteromeles arbutifolia (Lindley) Roemer +, RI Toyon; Christmas Berry Jun-Oct D Rosa californica Cham. & Schldl. +, RI, WI California Rose Spring-Summer M Rubiaceae — Madder Family *Galium aparine L. +, WI Comon Bedstraw; Goose Grass Mar-Jul M Salicaceae — Willow Family Populus fremontii S. Watson ssp .fremontii +, RI, WI Fremont or Alamo Cottonwood Spring M, I SalLx exigua Nutt. Sandbar; Narrow-LeavedWillow + Mar-May W, M Salix gooddingii C. Ball Goodding's Black Willow +, WI Mar-Apr W, M, S Salix laevigata Bebb Red willow IRWD Spring W, M Salix lasiolepis Benth. Arroyo Willow +, WI Feb-Apr M, I,S Saururaceae — Lizard Tail Family Anemopsis californica (Nutt.) Hook. & Am. +, WI YerbaMansa Mar-Sep W, M CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 63 Scrophulariaceae — Figwort Family Mimulus aurantiacus Curt. RI Bush Monkey Flower Spring-Late Spring *Veronica anagallis-aquatica L. +, WI Great Water Speedwell Europe May-Sep Veronica peregrina L. ssp. xalapensis (Kunth) Pennell Purslane sor Mexican speedwell Spring-Early Summer Solanaceae — Nightshade Family *Datura stramonium L. + Jimson Weed Apr-Sept Datura wrightii Regel + Jimson Weed Apr-Oct Lycium californicum Nutt. + California Box Thom Mar-Jul *Lycopersicon esculentum L. Tomato S. and C. Amer. Spring Nicotiana bigelovii (Torn.) Wats. var. wallacei Gray +, WI Wallace's Tobacco May-Oct *Nicotiana glauca Grah. +, WI Tree Tobacco S Am. Spr-Sum *Solanum americanum Mill. +, WI White Nightshade Mar-Sept Solanum douglasii Dunal in DC. +, WI Douglas' Nightshade Most mos. Tamaricaceae — Tamarisk Family *Tamarix chinensis Lour. +, WI Tamarix SWAsia Sum Urticaceae — Nettle Family Urtica dioica L. ssp. holosericea (Nutt.) Thome +, WI Hoary Nettle Jan-Apr * Urtica urens L. + Dwarf Nettle Europe Spring D S VPI, WI I M, I M, I, D D M S M, I, D I, D, S M, I, D, S W, M M, I M, I 64 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004) Monocotyledonf.s — Monocots Arecaceae (Palmae) — Palm Family *Phoenix canariensis Chabaud Canary Island Date Palm Canary Isl. ? M * Phoenix dactylifera L. + Date Palm N Africa ? M *Washingtonia robusta H. Wendl. + Mexican Fan Palm Baja, Mex. Early summer M, I, D Cyperaceae — Sedge Family Bolboschoenus maritimus (L.) Palla [<= Scirpus maritimus L.] +, WI Alkali Bulrush Spring s Cyperus eragrostis Lam. +, WI Tall Umbrella-Sedge Cyperus esculentus L. WI May-Nov W, M Yellow Umbrella-Sedge Cyperus odoratus L. WI Jun-Oct s Fragrant Umbrella-Sedge Eleocharis macrostachya Britton VPI, WI 7 w Creeping Spikerush Spring w Schoenoplectus americanus (Pers.) Volkart ex Schinz & R. Keller [<= Scirpus a. Pers.] +, WI Olney's Bulrush Jun-Aug Schoenoplectus californicus (C. A. Mey.) Sojak [<= Scirpus c. (C. Meyer) Steudel] California Bulrush Jun-Sep Schoenoplectus robustus (Pursh) M. T. Strong [<= Scirpus r. Pursh] Coastal Bulrush Apr-Aug Iridaceae — Iris Family Sisyrinchium be llum S. Wats. +, R1 Blue-Eyed Grass Spring D Juncaceae — Rush Family +, WI ? M, I VPI, WI Mar-Aug M Lemnaceae — Duckweed Family Lemna minuta Kunth +, WI Duckweed ? W Juncus balticus Willd. Wire Rush Juncus bufonius L. Toad Rush jS + jg + j* k^k^k CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 65 Liliaceae — Lily Family * Asparagus officinalis L. +, WI Garden Asparagus Europe Spring S Bloomeria crocea (Torrey) Cov. + Common Goldenstars Apr-Jun D Dichelostemma capitatum Alph. + Blue Dicks, Wild Hyacinth Mar-May I, D Chlorogalum pomeridianum (DC.) Kunth RI, + Soap Plant Apr-Jun D Poaceae (Gramineae) — Grass Family *Arundo donax L. +, WI Giant Reed Europe Mar- Sep M, S *Avena barbata Link + Slender Wild Oat Europe Mar-Jun S *Avena fatua L. + Wild Oat Europe Apr-Jun I, D *Avena sativa L. + Cultivated Oat Europe Spring D *Bromus diandrus Roth + Common Ripgut Grass Europe Mar-Jun I, D, S *Bromus hordeaceus L. + Soft Chess Eurasia Apr-Jul I, D, S *Bromus inermis Leyss. Smooth Brome Europe May-Aug I, D *Bromus madritensis L. ssp. rubens (L.) Husnot +, WI Foxtail Chess Europe Mar-Jun I, D *Cortaderia selloana (Schultes) Asch. & Graebner Pampas Grass S Am. Late Summer M *Crypsis schoenoides (L.) Lam. WI Swamp Grass Europe Jun-Jul I *Cynodon dactylon (L.) Pers. +, WI Bermuda Grass Jun-Aug M *Digitaria sanguinalis (L.) Scop. +, WI Crab Grass Europe Jun-Sep S Distichlis spicata (L.) E. Greene +, WI Coastal Salt Grass Apr-Jul M *Echinochloa colona (L.) Link + Jungle Rice Eurasia Jul-Oct M *Echinochloa crus-galli (L.) P. Beauv. +, WI Common Barnyard Grass Eurasia; Africa Jui-Oct S Hordeum brachyantherum Nevski ssp. californicum (Covas & Stebb.) Bothmer, N. Jacobsen & O. Seberg + California Barley Apr-Aug I, D *Hordeum murinum L. ssp. leporinum (Link) Arcangeli + Hare Barley Europe Apr-Jun I, D *Hordeum vulgare L. Cultivated Barley Europe Apr-Jul I, D 66 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] *Lamarckia aurea (L.) Moench + Goldentop Mediterranean Mar-Apr D Leymus condensatus (S. Presl) A. Love +, RI Giant Wild Rye Spring D Leymus triticoides (Buckley) Pilger +, WI Beardless Wild-Rye Jun-Jul I, D Leptochloa uninervia (C. Presl) A. Hitchc. & Chase +, W Dense-flowered Sprangletop Jul-Oct I *Lolium multiflorum Lam. + Italian Ryegrass Europe Jun-Aug I Nassella pulchra (A. Hitchc.) Barkworth + Purple Needlegrass Mar-May I, D *Paspalum dilatatum Poiret +, WI Dallis Grass S Am. May-Nov M Paspalum distichum L. IRWD Knot Grass Late Spring M *Pennisetum clandestinum Chiov. IRWD, WI Kikuyu Grass Africa Apr-Jun M, I *Phalaris minor Retz. + Little Seed Canary Grass Mediterranean Apr-Aug M, I, D *Phalaris canariensis L. + Canary Grass Medit. Eur. Apr-Aug M, I *Polypogon monspeliensis (L.) Desf. +, WI Annual Beard Grass Europe Apr-Aug M, I *Schismus barbatus (L.) Thell. + Mediterranean Schismus Eurasia; Africa Mar-Apr I *Setaria verticillata (L.) P. Beauv. +, WI Bur Bristlegrass Europe May-Sep ? *Sorghum halepense (L.) Pers. +, WI Johnson Grass Medit. Mar- Sep 7 *Vulpia mvuros (L.) K.C. Gmelin var. hirsuta (Hackel) Asch. & Graebner +, WI Foxtail Fescue Europe Mar-Jun I Potamogetonaceae — Pondweed Family Potamogeton pectinatus L. WI Fennel-leaved Pondweed Apr-Jun W Typhaceae — Cattail Family Typha angustifolia L. +, WI Narrow-Leaved Cattail Jun-Jul W, M Typha domingensis Pers. WI Southern Cattail Late Spring W, M Typha latifolia L. +, WI Broad-Leaved Cattail Jun-Jul W, M CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 67 NOTEWORTHY COLLECTION - CALIFORNIA Malacothrix saxatilis (Nuttall) Torrey & A, Gray var. saxatilis (Asteraceae) Discovered in Orange County, Southern California Richard E. Riefner, Jr. 5 Timbre, Rancho Santa Margarita, CA 92688 - and - Steve Boyd Herbarium, Rancho Santa Ana Botanic Garden 1500 North College Avenue, Claremont, CA 91711 Malacothrix saxatilis (Nuttall) Torrey & A. Gray var. saxatilis, cliff malacothrix, is a California Native Plant Society (CNPS) List 4 species, which is restricted to scrub habitats along the immediate coast only from Santa Barbara County and Ventura County in southern California (Davis 1993, Malacothrix, pp. 314-315, in Hickman (ed.), The Jepson Manual: Higher Plants of California, University of California Press Tibor (ed.) 2001, Inventory of Rare and Endangered Plants of California, Sixth Edition, California Native Plant Society, Sacramento). This perennial herb is a well-known component of the coastal bluff flora between Point Conception and Santa Barbara (Munz 1974, A Flora of Southern California, University of California Press; Abrams 1940, Illustrated Flora of the Pacific States, Stanford University Press), but it has not been reported from these floras or in treatments of the Asteraceae from local floras covering coastal Ventura, Los Angeles, Orange, and San Diego counties (Mattoni and Longcore 1997, Crossosoma 23:71-102; Wishner 1997, Flora of the Santa Monica Mountains: synonymized checklist and index, Crossosoma 23:3-63; Wishner 1998, Addendum, Crossosoma 24:111-112; Wishner 2000, Addendum II , Crossosoma 26:13-14; Wishner 2003, Addendum III, Crossosoma 28: 14; Roberts 1998, A Checklist of the Vascular Plants of Orange County, California, Second Edition, F.M. Roberts Publications, Encinitas; Simpson et al. 2002, Checklist of the Vascular Plants of San Diego County, California, San Diego Natural History Museum, www.sdnhm.org/research/botanv/sdplants/index.html Herein, we report the first known records of this taxon in Orange County, and describe its habitat and associated species. Orange Co., City of Laguna Beach, west of Pacific Coast Hwy, vicinity of Aliso Creek Park. Rare, growing in disturbed coastal bluff scrub dominated by exotic species of Acacia, Atriplex, Bromus, Cortaderia, Ipomoea, and Myoporum. Native species at the site include Dudleya lanceolata (Nutt.) Britton & Rose, Euphorbia misera Benth., Isomeris arborea Nutt., Encelia californica Nutt., Eriogonum fasciculatum Benth. var. foliolosum (Nutt.) Abrams, Isocoma menziesii (Hook. & Am.) G. Nesom, and Rhus integrifolia (Nutt.) Brewer & S. Watson. Location at 33.507°N 1 17.750°W, UTM Zone 1 IS N3707284 E430681 (NAD 27),elev<50 ft., USGS 7.5-minute quandrangle Dana Point, T8S R8W NW 1/4 of Section 5, 21 August \999, Riefner 99-360 (RSA); Orange Co., City of Dana Point, west of Pacific Coast Hwy, vicinity of Salt Creek Park. Locally abundant, growing in coastal bluff scrub dominated by native species including Atriplex californica Moq., A. lentiformis (Torrey) S. Watson ssp. 68 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004) lentiformis, Camissonia cheiranthifolia (Sprengel) Raim ssp. suffruticosa (S. Watson) Raven, Dudleya lanceolata, Isomeris arbor ea, Encelia californica, Eriogonum fasciculatum var. foliolosum, Lycium californicum Nutt., Rhus integrifolia, and Suaeda taxifolia (Standley) Standley. Non-native species invading the bluff site include species of Acacia , Atriplex , Bromus , Cortaderia, and Caprobrotus. Location at 33.481°N 117.724°W, UTM Zone 1 IS N3703885 B433056 (NAD 27), alt. <50 ft., U.S.G.S 7.5’ Quadrangle Dana Point, T8S R8W W 1/2 Section 16, 22 May 1999, Riefner 99-289 (RSA); same locality, 19 September 2000, Riefner 00-739 (RSA). Coastal bluff scrub is a community that is adapted to rocky or poorly developed soils exposed to salty, moisture-laden winds along the immediate coast (Holland 1986, Preliminary Description of the Terrestrial Natural Communities of California, Natural Heritage Program, California Department of Fish and Game, Sacramento). This community is highly vulnerable to extirpation by development and to degradation by exotic plants in the south coast region. For this reason, cliff malacothrix is included in the CNPS inventory of rare and endangered vascular plants with rarity- endangerment-distribution (RED) of 1-2-3, the “watch list.” It is also a California Department of Fish and Game “special plant” (California Department of Fish and Game, Natural Diversity Data Base, 2002, Special Vascular Plants, Bryophytes, and Lichens List, Sacramento). In southern Orange County, coastal bluff scrub is being rapidly replaced by residential development, irrigated ornamental plantings, exotic vegetation, and bluff stabilization projects, which often rely on traditional landscaping techniques that incorporate non-native, often highly invasive plants and concrete or rock rip-rap structures. A brief list of the aggressive exotic plants that are rapidly replacing native coastal bluff scrub species in south coastal Orange County includes acacia (Acacia longifolia (Andrews) Willd., giant reed ( Arundo donax L.), black mustard ( Brassica nigra [L.] Koch), brome grasses (Bromus spp.), pampas grass (Cortaderia selloana [Schultes] Asch. & Graebner), fennel (Foeniculum vulgare Miller), Perez’s sea-lavender (Limonium perezii [Stapf] Hubb.), myop (Myoporum laetum Forster), tree tobacco (Nicotiana glauca Graham), fountain grass (Pennisetum setaceum Forsskal.), Spanish sunflower (Pulicaria paludosa Link), castor bean (Ricinus communis L.), nasturtium (Tropaeolum majus L), several species of ice plants, Carpobrotus edulis (L.) N.E. Br., Malephora crocea (Jacq.) Schwantes, Mesembryanthemum crystallinum L., Tetragonia tetragonioides (Pallas) Kuntze, and several species of saltbush, most frequently, Australian saltbush (Atriplex semibaccata R. Br.). Cliff malacothrix grows with other CNPS List 4 plants in southern Orange County. At Laguna Beach, it is associated with Euphorbia misera, and in the Salt Creek area, north of Dana Point, it grows with Suaeda taxifolia and Lycium californicum. Each of these List 4 plants is threatened in south coastal Orange County. If they are to persist in the wild, these plants will require careful monitoring and protection from habitat degradation by exotic species and habitat loss owing to unmitigated development. There has been little disagreement regarding the circumscription of the majority of species in the genus Malacothrix, with the exception of the annual species endemic to the California Islands (Davis 1997, Madrono 44:223-244). Accordingly, the varieties of perennial Malacothrix are easily separated. Malacothrix saxatilis var. saxatilis is easily distinguished from the more common and widespread var. tenuifolia (Nutt.) A. Gray by its generally linear to ovate, entire upper leaves, and stems that are leafy to the inflorescence (see the illustration in Abrams 1940, loc. cit.). In Orange County, M. s. var. saxatilis has a leathery texture, large conspicuous involucres, and generally it flowers later in the season than coastal populations of the var. tenuifolia. We found that M. s. var. CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 69 saxatilis blooms most frequendy in late August and/or in September, but may flower in early spring during wet years. Another plant that is most easily detected during late summer surveys, Baccharis malibuensis R. M. Beauch. & Henr. (CNPS List IB), was recently discovered in Orange County (Boyd 2002, Madrono 49:54); it was previously known only from extreme western Los Angeles County. Therefore, it is not completely surprising to find another late blooming, more northern plant such as cliff malacothrix in Orange County. Malacothrix saxatilis var. saxatilis should be sought at additional localities in coastal southern California, including the Channel Islands, and on the mainland from cliff and scrub habitats at Point Sherman Park and at the Palos Verdes Peninsula in Los Angeles County, at Abalone Point and San Clemente in Orange County, and on the bluffs at San Onofre in San Diego County. We recommend that in late August and September focused surveys should be conducted as standard protocol, in order to thoroughly document the occurrence of all special-status species that would otherwise not be found during traditional spring to early summer floristic survey programs. 70 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Book Review COAST REDWOOD: A NATURAL AND CULTURAL HISTORY Edited by John Evarts and Marjorie Popper Written by Michael Barbour, Sandy Lydon, Mark Borchert, Marjorie Popper, Valerie Whitworth, and John Evarts Published in 2001 by Cachuma Press, P.O. Box 560, Los Olivos, California Specifications: 228 pages, 230 color and 100 black and white illustrations, index. Price: ISBN 0-9628505-5-1, Paperback, $27.95; ISBN 0-9628505-6-x, Hardcover, $37.95. Some of you will remember my glowing reviews of two other books from Cachuma Press, namely. Oaks of California and Conifers of California. Well, this book about Coast Redwood follows the honored tradition. Evarts and Popper exhibit a fine talent for mixing botanical and cultural information with copious illustrations and photographs. Other books that feature giant trees have been published recently, but this volume celebrates a single species, Sequoia sempervirens (D. Don) Endl., the tallest trees in the world, in a way that is unrivaled in publishing. Coast Redwood begins with a chapter on the origin and distribution of Coast Redwoods. Here, the history of the discovery of these trees is outlined as early explorers such as Gaspar de Portola and early botanists such as Thaddeus Haenke raved about these marvelous trees. The original name, Palo Colorado (Red wood), is attributed to father Juan Crespi, a Franciscan missionary who accompanied Gaspar de Portola, and described the trees from near the Pajaro River in the Santa Cruz Mountains. The genetics and evolution of the species is also described in this chapter. Further chapters include: Life of a Coast Redwood, Ecology of the Coast redwood Forest, Wildlife of the Coast redwood forest. Harvest and Utilization of Coast Redwood, History of Coast Redwood Preservation, and finally, Conservation and Management of Coast Redwood Forests. The book concludes with a series of appendices that summarize comparisons of other species of redwoods, average temperature and precipitation records, common and scientific names of plant species that grow with redwoods, timber harvests, and Coast Redwood conservation organizations. The life history and ecology of Coast Redwoods encompasses less than half of the total book, but that does not lessen its value to biologists. The parts that are there are well done, including the sections that illustrate and describe the animals of the redwood forests. The ecological sections are written so that lay persons can understand them, but they are not so “dumbed-down” that they lack relevance for biologists. Some lesser-known species dependent upon redwood forests, such as the Marbled Murrelet, are discussed thoroughly in sidebars, whereas, other better-known animals such as the Roosevelt Elk, considered by some authors to be the wildlife symbol of the redwood forest, are only covered in a cursory way. Scattered throughout the book there also are sidebars that feature certain specific points of interest about Coast Redwoods. For example, one sidebar entitled “To find the Biggest Tree” features the history of documenting the tallest trees from “El Palo Alto” at 137 feet in 1776, to the current “Stratosphere Giant” at 386.6 feet tall. Another sidebar explains why California has two state trees, both redwoods. The “California Redwood” was named the official state tree in 1931 by the state legislature, but no scientific name was assigned. In 1951, Coast Redwood and Sierra Redwood (both in the genus Sequoia at the time) were established as the state tree, and those common names also were made official by the State Parks Commission and the State Forester. CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 71 It is interesting that these magnificent trees have not always inspired reverence and awe. It has been pointed out that for many years, loggers have viewed the trees as a commodity. The story of Julia Butterfly Hill, who set up camp for two years in the top of a 1,000-year-old redwood named “Luna,” is documented in another sidebar. Her protest against the actions of Pacific Lumber Company coincided with the contentious negotiations that ultimately led to the purchase of the Headwaters Forest, wherein 5,600 acres of redwood forest land were transferred to federal ownership. Not only is the text well done, but the photographs are excellent, ranging from historic black and whites, to fine color images. All of this is enhanced by high quality glossy paper that enables the photographs to be published in crisp detail. Kudos to the fine team of researchers who located and reprinted the historical photographs. If you are a naturalist and you love books, don’t hesitate to purchase this volume for your library. — Allan A. Schoenherr is Professor of Ecology at Fullerton College, — author of A Natural History of California, — and lead author of Natural History of the Islands of California — ( both from UC Press) 72 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004) INDEX AND BIBLIOGRAPHY OF CROSSOSOMA VOLUME 29, 2003 Crossosoma, the Journal of the Southern California Botanists, Inc., starting in May 1975 as SCB News , is now in the 29th year of publication. A comprehensive index and bibliography for the first 26 years of the journal appeared in Crossosoma 26(2). Indexes and bibliographies for subsequent volumes appear in Crossosoma 27(2) and Crossosoma 28(2). This and previous indexes contain at least two entries, and frequently several more than that. At the minimum, authors and co-authors are indexed, and at least one key word from the title. Articles dealing with various aspects of a particular species' biology are indexed both by species name and family, and occasionally by common name, and also by recent nomenclature changes (i.e. Stipa changed to Nassella and Achnatherum) reflected in The Jepson Manual or subsequently posted at the Jepson Online Interchange website. Species names that are indexed are also given a brief descriptive phrase to indicate the nature of the subject, for example: "Stromatella bermudana, noteworthy collection of, Riverside Co.," and more. Where articles pertain to specific locations, these locations are indexed, along with a descriptive phrase. Where such locations are in California, the county or counties are also indexed. A few other recurring items such as book reports, and annual symposium programs are also tracked herein. In all, approximately 148 articles, editorials, photographic and artistic submissions have appeared in Crossosoma, dealing with diverse aspects of botany in southern California, Baja California, and elsewhere. A bibliography of articles that appear in the current volume follows the index. These index and bibliographic additions are being appended to the cumulative index, in anticipation of future website availability. Some people might ask “What is the benefit of including the current index in the current volume?” The answer is quite simple. Early compilation assures that an index for the volume gets prepared, and the procedure does not rely on an uncertain future effort to compile and publish one. A timely compilation of the index and bibliography is very important for the continuity and long-term value of the journal. — Carl Wishner, Editor, Compiler CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 73 careful monitoring and protection from habitat degradation by exotic species and habitat loss owing to unmitigated development. There has been little disagreement regarding the circumscription of the majority of species in the genus Malacothrix. with the exception of the annual species endemic to the California Islands (Davis 1997, Madrono 44:223-244). Accordingly, the varieties of perennial Malacothrix are easily separated. Malacothrix saxatilis var. saxatilis is easily distinguished from the more common and widespread var. tenuifolia (Nutt.) A. Gray by its generally linear to ovate, entire upper leaves, and stems that are leafy to the inflorescence (see the illustration in Abrams 1940, loc. cit.). In Orange County, M. s. var. saxatilis has a leathery texture, large conspicuous involucres, and generally it flowers later in the season than coastal populations of the var. tenuifolia. We found that M. s. var. saxatilis blooms most frequently in late August and/or in September, but may flower in early spring during wet years. Another plant that is most easily detected during late summer surveys, Baccharis malibuensis R. M. Beauch. & Henr. (CNPS List IB), was recently discovered in Orange County (Boyd 2002, Madrono 49:54); it was previously known only from extreme western Los Angeles County. Therefore, it is not completely surprising to find another late blooming, more northern plant such as cliff malacothrix in Orange County. Malacothrix saxatilis var. saxatilis should be sought at additional localities in coastal southern California, including the Channel Islands, and on the mainland from cliff and scrub habitats at Point Sherman Park and at the Palos Verdes Peninsula in Los Angeles County, at Abalone Point and San Clemente in Orange County, and on the bluffs at San Onofre in San Diego County. We recommend that in late August and September focused surveys should be conducted as standard protocol, in order to thoroughly document the occurrence of all special-status species that would otherwise not be found during traditional spring to early summer floristic survey programs. 74 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Fraga, Naomi Crossosoma 29(2):4 1 -43 index and bibliography to Crossosoma : Volume 29 2003 Crossosoma 29(2):72-76 Inyo National Forest: progress report toward a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):4 1 -43 ; photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2):44 Irvine, University of California, San Joaquin Freshwater Marsh Reserve, Orange County, vascular plant checklist of Crossosoma 29(2):45-66 Kern Co.: Sierra Nevada, progress report toward a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):41-43; photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2):44 Knudsen, Kerry Crossosoma 29(l):35-38 Laguna Beach, City of: Malacothrix saxatilis var. saxatilis noteworthy collection of Crossosoma 29(2):67-69 lichens: noteworthy collections Crossosoma 29(l):35-38; enhance recruitment success of rare Dudleya species Crossosoma 29(1): 1-34 Lichinaceae: Stromatella bermudana noteworthy collection of, Riverside Co., San Bernardino National Forest, San Jacinto Mountains, East Canyon at end of Morris Ranch Road Crossosoma 29(l):35-38 Los Angeles Co.: Santa Monica Mountains: Agoura Hills; Texosporium sancti-jacobi noteworthy collection of Crossosoma 29(l):35-38 Malacothrix saxatilis var. saxatilis noteworthy collection of, City of Laguna Beach, west of Pacific Coast Hwy, vicinity of Aliso Creek Park Crossosoma 29(2):67-69 Mojave Desert: progress report toward a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):41 -43; photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2) :44 Mulroy, ThomasW. Crossosoma 29(1): 1-34 Niebla ceruchoides (the pincushion lichen) enhance recruitment success of rare Dudleya species Crossosoma 29(1): 1-34 noteworthy collections: lichens Crossosoma 29(l):35-38; Malacothrix saxatilis var. saxatilis in Orange Co. Crossosoma 29(2):67-69 Orange Co.: Malacothrix saxatilis var. saxatilis noteworthy collection of. City of Laguna Beach, west of Pacific Coast Hwy, vicinity of Aliso Creek Park Crossosoma 29(2):67-69; University of California Natural Reserve System, San Joaquin Freshwater Marsh Reserve, Irvine, Orange County, vascular plant checklist of Crossosoma 29(2):45-66 Owens Peak Wilderness progress report tow ard a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):41-43; photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2):44 photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2):44 phytogeography; of lichens in southern California Crossosoma 29(l):35-38; of Sierra Nevada and Mojave Desert Crossosoma 29(2):41-43 pincushion lichen see Niebla ceruchoides progress reports: Toward a flora of the Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):4 1 -43 Ramalinaceae: Niebla ceruchoides (the pincushion lichen) enhance recruitment success of rare Dudleya species Crossosoma 29( 1): 1 -34 rare plants: recruitment success of rare Dudleya species enhanced by lichens Crossosoma 29(1): 1-34 redwood, coast see Sequoia sempervirens Riefner, Richard E. Jr. Crossosoma 29(1): 1-34; Crossosoma 29(2):67-69 Riverside Co.: Stromatella bermudana noteworthy collection of, San Mateo Wilderness Area, San Mateo Canyon Crossosoma 29(1 ):35-38; Catapyrenium squamellum noteworthy CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] 75 collection of, San Bernardino National Forest, San Jacinto Mountains, East Canyon at end of Morris Ranch Road Crossosoma 29( 1 ):35-38 San Bernardino National Forest: Stromatella bermudana noteworthy collection of, San Mateo Canyon, Riverside Co. Crossosoma 29(l):35-38 San Jacinto Mountains: Stromatella bermudana noteworthy collection of, Crossosoma 29(l):35-38 San Joaquin Freshwater Marsh Reserve, University of California Natural Reserve System, Irvine, Orange County, vascular plant checklist of Crossosoma 29(2):45-66 San Mateo Canyon, Riverside Co. Stromatella bermudana noteworthy collection of Crossosoma 29(l):35-38 San Mateo Wilderness Area, Riverside Co., Stromatella bermudana noteworthy collection of Crossosoma 29(l):35-38 Santa Monica Mountains: Texosporium sancti-jacobi noteworthy collection of, Cornell Corners, Los Angeles Co. Crossosoma 29(l):35-38 SCB Source and Use of Funds 2003 Crossosoma 29(l):39-40 SCB Grant Program: progress report toward a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):41-43 Schoenherr, Allan A. Crossosoma 29(2):70-71 Sequoia sempervirens, book entitled Coast Redwood (Barbour et al.) is reviewed by Allan A. Schoenherr Crossosoma 29(2):70-71 Sierra Nevada: progress report toward a flora of Owens Peak eastern watershed, Kern Co. Crossosoma 29(2):4 1 -43; photograph: challenging terrain of the Owens Peak eastern watershed Crossosoma 29(2):44 soil crusts: enhance recruitment success of rare Dudleya species Crossosoma 29( 1 ): 1 -34 Source and Use of Funds 2003 Crossosoma 29(l):39-40 Southern California Botanists, Inc.: SCB Source and Use of Funds 2003 Crossosoma 29(l):39-40 Stromatella bermudana noteworthy collection of, Riverside Co., San Bernardino National Forest, San Jacinto Mountains, East Canyon at end of Morris Ranch Road Crossosoma 29(l):35-38 symposia, programs: (note: program abstracts not published in Crossosoma after 1991, but continuing to present in Leaflets ); Back From the Brink: Conservation Success Stories [2003 Oct 18] Taxodiaceae see Sequoia sempervirens Texosporium sancti-jacobi noteworthy collection of, Los Angeles County, Santa Monica Mountains, Agoura Hills, Cornell Corners Crossosoma 29(l):35-38 University of California: Natural Reserve System, San Joaquin Freshwater Marsh Reserve, Irvine, Orange County, vascular plant checklist of Crossosoma 29(2):45-66 Verrucariaceae: Catapyrenium squamellum noteworthy collection of. Riverside Co., San Mateo Wilderness Area, San Mateo Canyon Crossosoma 29(l):35-38 wilderness area: San Mateo, Stromatella bermudana noteworthy collection of, San Mateo Canyon, Riverside Co. Crossosoma 29(l):35-38 Wishner, Carl Crossosoma 29(1): 1-34; Crossosoma 29(2):72-76; Crossosoma 29(2):44 76 CROSSOSOMA 29(2), Fall-Winter 2003 [issued August 2004] Bibliography of Articles Published in Crossosoma Volume 29, 2003 Bowler, Peter A„ and Mark A. Elvin. 2003 [issued July 2004]. The Vascular Plant Checklist for the University of California Natural Reserve System’s San Joaquin Freshwater Marsh Reserve. Crossosoma 29(2):45-66. Fraga, Naomi. 2003 [issued July 2004]. The vascular flora of the Owens Peak eastern watershed: A review of progress [Sierra Nevada, Kern Co., California], Crossosoma 29(2):4 1 -43. Knudsen, Kerry. 2003 [issued July 2004]. Three notable lichen collections and their relationship to lichen distributions in Southern California. Crossosoma 29(l):35-38. Riefner, Richard E. Jr., and Steve Boyd. 2003 [issued July 2004], Malacothrix saxatilis (Nuttall) Torrey & A. Gray var. saxatilis (Asteraceae) discovered in Orange County, Southern California. Crossosoma 29(2):67-69. Riefner, Richard E. Jr., Peter A. Bowler, Thomas W. Mulroy, and Carl Wishner. 2003 [issued July 2004]. Lichens on rock and biological crusts enhance recruitment success of rare Dudleya species (Crassulaceae) in Southern California. Crossosoma 29(1): 1-34. Schoenherr, Allan A. (reviewer). 2003 [issued July 2004], Book Review — Coast Redwood: A Natural and Cultural History, By M. Barbour et al. 2001 [Cachuma Press] Crossosoma 29(2):70-71. Wishner, C. (compiler). 2003 [issued July 2004]. Index and bibliography of Crossosoma Volume 29, 2003. Crossosoma 29(2):72-76. Wishner, C. (photographer). 2003 [issued July 2004], Photograph: Challenging terrain of the Owens Peak eastern watershed. Crossosoma 29(2):44. Crossosoma Volumes and Issues Indexed The first four issues of the journal, comprising volume 1 and the first issue of volume 2 were called SCB News. Starting in May 1976, the journal was titled Crossosoma. Volume numbers and issue numbers did not appear explicitly on the covers of the journal until volume 5 number 1 (1979). In some cases over the years, erroneous volume numbers, issue numbers, or dates were printed on the original covers, and some issues were undated. See Crossosoma 26(2):29-52 for a comprehensive index and bibliography of Volumes 1 through 26, 1975-2000. The index and bibliography for Volume 27 [2001] appears in Crossosoma 27(2):58-60. The index and bibliography for Volume 28 [2002] appears in Crossosoma 28(2):50-54. Vol. 29 2003 Spring-Summer 29(1) [issued July 2004] Fall-Winter 29(2) [issued July 2004] 3 5185 00267 9858 Southern California Botanists , Inc. — Founded 1927 — Memberships, Subscriptions, Back Issues Individual and Family Memberships in SCB are $15 per calendar year domestic, and $20 per year to foreign addresses. Membership includes two issues of CROSSOSOMA, and 5 or 6 issues of Leaflets , the newsletter of SCB. Leaflets provides time-dated information on activities and events that may be of interest to our membership. A subscription to CROSSOSOMA is available to libraries and institutions at the domestic rate of $25 per calendar year, and $30 to foreign institutions. Back issues (Volumes 18-present) are available for $5 each, or $10 for the volume, postpaid. Prior to Volume 18, CROSSOSOMA included time-dated notices to the membership and was published six times a year. These back issues of Volumes 1-17 are $1 each, or $6 per volume, postpaid. Some back issues that are out of stock may be provided as photocopies. SCB Special Publications No. 1 A Flora of the Santa Rosa Plateau, by Earl W. Lathrop and Robert F. Thome, 39 pp $7.00 No. 3 Endangered Plant Communities of Southern California, Proceedings of the 15th Annual SCB Symposium, edited by Allan A. Schoenherr, 1 14 pp $12.00 [Special Publication No. 2, Flora of the Santa Monica Mountains, 2nd ed., by Peter H. Raven, Henry J. Thompson, and Barry A. Prigge is out of print] Book prices include California state sales tax, handling, and domestic postage. By request, the following article has been reprinted as a separate, with covers, and is available for plant collecting workshops: Reprint. Herbarium Specimens as Documents: Purposes and General Collecting Techniques, by T. S. Ross [from CROSSOSOMA 22(1 ):3-39, 1996] $3.95 each; 10 for $22.50 Applications for membership, requests for purchases of Special Publications and back issues, name or address corrections, and requests for replacement of lost or damaged CROSSOSOMA issues should be sent to Alan Romspert, Treasurer Southern California Botanists, Inc., Department of Biology, California State University, Fullerton, CA 92834, USA. Consult the website for current email contact information. Make your check or money order payable to Southern California Botanists, or SCB. http://socalbot.org LUESTERT. MERTZ LIBRARY JUN 2 3 2005 NEW YORK BOTANICAL GARDEN