CROSSOSOMA Journal of the Southern California Botanists, Inc. Cryptantha of Southern California Volume 35, Number 1 Spring-Summer 2009 Southern California Botanists, Inc. -Founded 1927 - http://www. socalbot. org CROSSOSOMA (ISSN 0891-9100) is published twice a year by Southern Cali- fornia Botanists, Inc., a California nonprofit organization of individuals devoted to the study, conservation, and preservation of the native plants and plant com- munities of southern California. SCB Board of Directors for 2009 President Gary Wallace Vice President Naomi Fraga Secretary. Carrie Kiel Treasurer Alan P. Romspert Webmaster Naomi Fraga Editors of Crossosoma Scott D. White and Michael Honer Editor of Leaflets Kerry Myers Directors-at-large David Bramblet Gilberto Ocampo Sara Baguskus Linda Prince Elizabeth Delk Gina Richmond Erica Gardner Fred Roberts Charlie Hohn Darren Sandquist Diane Menuz Allan Schoenherr Orlando Mistretta Jonathan Snapp-Cook Bart O’Brien Sula Vanderplank and Katie Vinzant Ex offidio Board Member Sula Vanderplank (Past President) Articles, book reviews, or other items for submission to CROSSOSOMA can be sent to the editor Scott White (scottbioservices@verizon.net) or 201 N. First Ave., #102, Upland, CA., USA, 91786. Electronic submission is preferred. Please see our website, www. socalbot.org, for format guidelines. Notices of a time-dated nature (field trips, work- shops, symposia, etc.) to be included in the newsletter Leaflets should be submitted to Kerry Myers, Editor of Leaflets, kerrymyers@fs.fed.us, or mail to: Kerry Myers, Botanist, SBNF Mountaintop Ranger Dist., 42300 North Shore Dr., Fawnskin, CA 92333-04004. 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 specifically stated. Copyright © 2009 by Southern California Botanists, Inc. All rights reserved. Permission to reproduce items in CROSSOSOMA, in whole or part, should be requested from the Editor. Crossosoma 35(1). Spring-Summer 2009 1 CRYPTANTHA OF SOUTHERN CALIFORNIA Michael G. Simpson and Kristen E. Hasenstab Department of Biology San Diego State University San Diego, California 92 182 USA msimpson@sunstroke.sdsu.edu: kristen.hasenstab@cgu.edu (Current address for K. Hasenstab: Rancho Santa Ana Botanic Garden. Claremont, 1 500 N. College Ave., California 91711) ABSTRACT: The genus Cnptantha (Boraginaceae) contains 202 species, with 49 species and 56 taxa (including varieties) occurring in Southern California, defined here as including the entire Southwestern California region and Tehachapi Mountain region of the California Floristic province, the entire Desert province, and most of the White and Inyo Mountain subregion of the Great Basin province. The purposes of this article are 1) to summarize the current taxonomy of Cryptantha species and infraspecies in Southern California: 2) to provide taxonomic keys and images illustrating the diagnostic features for identification: and 3) to review the distribution, environmental factors, and current conserv ation status of these taxa. KEYWORDS: Cryptantha , Boraginaceae. taxonomy. INTRODUCTION Taxonomic History and Nomenclature Cryptantha Lehmann ex G. Don, commonly known eye." is a genus within the family Boraginaceae. The has changed repeatedly over the last twenty years [Engler and Prantl 1897. Hey wood et al. 2007, Gottschling et al. 2001. Angiosperm Phylogeny Group (APG II) 2003]. with various authors recognizing either a broad or narrow family concept. Here we accept the APG II (2003) system of classification, which recognizes a broad Boraginaceae. As treated in this manner, the family may be divided into subfamilies Boraginoideae, Cordioideae. Eretioideae. Heliotropoideae. Hydrophylloideae, and (possibly) Lennoideae (see Stevens 2001 onwards). Despite these changes in classification. Cnptantha firmly belongs in subfamily Boraginoideae. characterized by an inflorescence that is a scorpioid cyme (Buys and Hilger 2003). a deeply 4-lobed ovary w ith a gynobasic style, and a fruit that is a schizocarp of nutlets. Cryptantha is an amphitropicallv distributed genus, with taxa in both western North America and western South America, but lacking populations in between. The genus is currently catalogued at 202 species. The greatest species diversity. 136 species, occurs in western North America, with distributions from Alaska to southern Mexico, and ifentificatK% NUV 9 VrtW NEWYO.Kk afc "pot circumscription of this family 2 Crossosoma 35(1), Spring-Summer 2009 as far east as Texas (Johnston 1925; Payson 1927; Higgins 1971; Mabberley 2008). Approximately 69 species occur in western South America, in Peru. Chile, Bolivia, and east to Argentina (Reiche 1915; Johnston 1927; Schwarzer 2007; Zuloaga et al. in prep). Three species, C. albida , C. circumscissa. and C. maritima , are distributed in both North and South America. Members of the genus are annual, biennial, or perennial herbs. Stems are simple to highly branched, generally ascending to erect, and densely covered in trichomes (Johnston 1925; Payson 1927; Munz and Keck 1959, 1968; Higgins 1971; W.A. Kelley and Wilken 1993, Mabberley 2008; R.B. Kelley in prep). Often in perennial taxa, a basal rosette of robust leaves is present before and during inflorescence growth. Among annuals, the basal rosette is common early in development but is lost as the intemodes of the aerial shoots (including inflorescence) elongate. Cauline leaves range from opposite to alternate in arrangement. Leaf shape is spatulate. lanceolate, oblanceolate, but most often linear. Leaves have acute to obtuse bases. Leaf vestiture is strigose, "rough hairy,” or "bristled.” The largest, relatively stout trichomes, which are often designated as bristles, often have bulbous bases, described as pustulate (giving a tessellate appearance to the leaf surface). These pustules are composed of epidermal cells arranged in a ring around the trichome base; the epidermal cells are elevated, opaque, and silicified and are thought to be cystolith in origin (Johnston 1925). The trichomes are most often translucent and white, but can be yellow-brown in some taxa. The inflorescence is characteristic of subfamily Boraginoideae, being a circinate (coiled), scorpioid cyme unit (often described as a helicoid cyme, but see Buys and Hilger 2003); however, the inflorescence structure is often obscured by short intemodes and absence of floral bracts. The calyx, considered part of the fruiting unit, is accrescent, and continues to grow' as the nutlets mature. The shape and fusion of the calyx, as well as calyx vestiture and orientation when the plant is in fruit, varies among taxa and is often used to distinguish them. Flowers are chasmogamous (opening and capable of cross-pollination) in most taxa, but may be cleistogamous (not opening and self-pollinated) in members of subgenera Cryptantha and Geocatya (modified into lenticular structures in the last). The corolla limb (the expanded portion above the corolla tube, containing the five lobes) ranges in size from less than 1 mm wide (measured across from lobe to lobe) to up to 2 cm wide. Corollas are almost universally white (a few species having yellow’ corollas), and are rotate to salverform. The corolla tube is usually equal to the calyx in length and bears five epipetalous stamens. At the outer corolla throat, the tissue is invaginated, resulting in five pinched or folded regions, known as fornices. The fornices, which enclose the throat, can vary in size, shape, and color, but are often yellow and possibly function as nectar guides. Corolla lobes are ovate-oblong or suborbicular, ranging from widely spreading to ascending in chasmogamous species. As is characteristic of the subfamily Boraginoideae, the ovary is deeply four-lobed. with a gynobasic style arising from the base and center of the ovary lobes. The ovary lobes mature into 1-seeded units with hardened pericarps, termed nutlets. The number of nutlets that develop to maturity can vary from 1 to 4. The tissue between the ovary lobes, interpreted as either receptacular tissue or the style base, continues to elongate and differentiate during fruit maturation. This tissue Crossosoma 35( 1 ), Spring-Summer 2009 3 is often called the gynobase and is that to which the nutlets are laterally attached at maturity. The gynobase is generally narrow ly-pyramidal in form, elongating during fruit maturation. (Note that “fruit’ ' is used here to include the nutlets and accompany ing calyx.) Nutlets are generally ovate to lanceolate in shape. The fruits of annual taxa are generally deciduous, and the whole unit encased by the calyx will easily detach from the inflorescence; however, in the perennial subgenus Oreocarya . the nutlets detach easily but the calyx and pedicel remain firmly attached to the plant. Several annual species in North American have fruits comprised of heteromorphic nutlets. Often, there are three smaller, easily detached nutlets, and one larger nutlet that is strongly adnate to the gynobase. The pericarp wall is variable in sculpturing and color. The attachment scar is generally a shallow, triangular areole (generally not rimmed or elevated), which is continuous with the ventral groove. When describing the ventral groove, morphologists generally consider the groove and attachment scar as one feature on the nutlet; thus the ventral groove is often described as being basallv forked. The genus name Cryptantha was first used by Johann Georg Christian Lehmann in a seed catalogue for the Hamburg Botanical Garden. In 1836, Fischer and Meyer used Lehmann's proposed name Cryptantha in association w ith two South American species. C. glomerata and C. microcapa. but with no formal genus diagnosis. In that same year, how ever, the name Cryptantha was validly published in the General System of Gardening and Botany by Don (1836), w ho provided the first formal diagnosis for the genus in association with the same two South American species (Johnston 1925). Thus, authorship of the genus should correctly be cited as Cryptantha Lehmann ex G. Don. Greene (1887a,b,c) was the first botanist to carefully study the borages of tribe Eritrichieae (within which Cryptantha has been classified), which are diverse in western North America. He authored several revisions of Eritricheae members, including Cryptantha . in a series of papers in the journal Pittonia. Green recognized the importance of classifying the group based on characters other than floral appearance, noticing the great variation in nutlet characteristics among members of this tribe. He expanded the genus Cryptantha to include North American species that had been placed in the genus Krynitzkia Fisch. & C.A. Mey. Greene’s circumscription of Cryptantha included strictly annual taxa with deciduous fruits. Additionally, Greene recognized several genera that are now treated as part of Cryptantha , including Oreocarya Greene. Piptocalyx Torr. [=Gree neochar is Giirke & Harms], and Eremocatya Greene. The next major revision came in the dissertation research of the Boraginaceae expert I. M. Johnston in 1925. In this monographic treatment of the North American species of Cryptantha. the genus was circumscribed to include Eremocary'a and Piptocalyx [=Greeneocharis]. Each species was revised and clarified in this work, with each of the 57 species placed into one of fifteen sections. Shortly after the publication of his dissertation. Johnston spent several months in South America. These studies, w ith new herbarium vouchers, led Johnston to conclude that the perennial Oreocarya species should also be included in Cryptantha to form one large, homogenous genus. In 1927, Johnston published treatments of South American Boraginaceae. recognizing Cryptantha as having three subgenera in South America: Krynitzkia . Geocarya . and Eucryptantha [ =Cryptantha ]. Also in 1927, Pavson. who was working on a 4 C-rossosoma 35(1), Spring-Summer 2009 Table 1. Subgenera of Ciyptantha recognized by Johnston (1927) and Payson (1927). Subgenus Characters delineating subgenus Krynrtzkia (incl. Eremocaryci and Greene ochar is) Oreocarya Geocarya Eucryptantha [=Crvptantha\ Plants with cleistogamous flowers. Cleistogamous flowers similar to chasmogamic flowers, except for closed corolla. Cleistogamous flowers located in axiles of leaves and often throughout the inflorescence. Distributed in South America. Plants with cleistogamous flowers. Basal cleistogamous flowers highly specialized into lenticular structures. Distributed in South America. Annual plants having only chasmogamous flowers. Distributed in North and South America. Perennial or biennial herbs, producing only chasmogamous flowers. Restricted to North America. monograph of Oreocaiya, agreed with Johnston that Oreocarya should be included in Ciyptantha. Payson, following Johnston's treatments, recognized four subgenera in Cryptantha : Oreocarya, Kiynitzkia, Geocarya and Eucryptantha [=Cryptantha\. (See Table 1 for details of the characters used to circumscribe these subgenera, and their distributions.) Since the 1920s, systematists have made revisions among species and their relationships within Cryptantha, but the subgenera themselves have remained largely unchanged. Higgins, another expert on the perennial taxa, published a revised monograph of Oreocarya in 1971, and agreed with Johnston and Payson on the inclusion of Oreocarya within Cryptantha. However, the classification of Oreocarya has been contentious. For example, Abrams (1951) recognized Oreocarya as genus separate from Cryptantha in his Illustrated Flora of the Pacific States: Washington, Oregon and California. More recently, Weber (1987) also recognized Oreocarya at the genus level, giving no specific reason for his circumscription, but arguing that genera should consist only of species easily recognizable by suites of characters. However, most recent classifications follow the treatments of Johnston, Payson, and Higgins in circumscribing Cryptantha in the broad sense, as encompassing all four subgenera. For Southern California, these include the treatments of Cryptantha in Munz (1974), The Jepson Manual (W.A. Kelley and Wilken 1993), and the draft treatment for the second edition of The Jepson Manual (R.B. Kelley in prep). Hasenstab (2009), in a molecular phylogenetic analysis of Cryptantha and relatives, concluded that the genus Cryptantha as currently defined appears to be paraphyletic, warranting the resurrection of several genera, including Eremocarya, Greeneocharis, and Oreocarya. Although we feel it premature to divide the genus for this article, the key we have prepared reflects (in part) a more natural grouping of taxa. Definition of the Southern California Region As this article is on the Cryptantha of Southern California, we must define what we mean by that geographic designation. We use Munz’s concept of Southern California, as he recognized it in A Flora of Southern California (1974). He defined the northern boundary of Southern California as "Point Conception, eastward along the crest of Crossosoma 35( 1 ). Spring-Summer 2009 5 > White & Inyo Mtns, Mr. Tehachapi fJV/, Mountains Desert Province Ventura Co. Los Angeles Co. Pt. Conception Southwestern Region Riverside Co. Imperial Co. Figure 1. Regions and provinces from The Jepson Manual (Hickman 1993). corresponding to Southern California, sensu Munz (1974). See this work for included subregions and districts. Ct*t. 6 Crossosoma 35(1), Spring-Summer 2009 the Santa Ynez mountains to the Mt. Pinos region in Ventura County, Fort Tejon in Kern County, the Tehachipi and Piute mountains, then northward to Little Lake in Inyo County and along the east slopes of the Inyo and White mountains to the Deep Springs region.” Munz stated that the region does not include the northern part of Santa Barbara County, the southern end of the San Joaquin Valley, nor any of the Owens Valley, as these regions contain more of a northern element of the flora (Figure 1). This boundary for '"Southern California” contains components of three floristic provinces designated in The Jepson Manual (Hickman 1993): 1 ) The entire Southwestern California region (including the South Coast, Channel Islands, Transverse Ranges, and Peninsular Ranges subregions) and the Tehachapi Mountain subregion of the California Floristic province; 2) the entire Desert province (Sonoran and Mohave deserts, including the Desert mountains); and 3) the White and Inyo Mountain subregion of the Great Basin province. Although ‘"Southern California” is often viewed as a socio- political designation, the region as defined here actually corresponds to major floristic provinces, regions, or subregions of California and thus has some scientific meaning. With regard to county borders. Southern California includes all of San Diego, Imperial, Los Angeles, Orange, Riverside, Ventura, and San Bernardino counties, the extreme southern and southeastern portion of Santa Barbara County, the extreme southern, south-central, and eastern parts of Kern County, all except the extreme northwestern and a small northeastern part of Inyo County, and a southeastern swath of Mono county (Figure 1). Identification of Southern California Cryptantha C.iyptantha species and infraspecies are generally recognized as being difficult to identify, even by professional botanists; the taxa are often being perceived to look very similar to one another. Perhaps the major reason for this perception is that the main feature used in identifying flowering plants - the flower itself - is generally both obscure and uniform among Cryptantha taxa. The corolla, the primary cue of identification both for humans and potential pollinators, is rarely used in Cryptantha identification, except for some variation in size and (rarely) in color. And yet, it should be pointed out that Cryptantha taxa vary tremendously in a great number of features, only some of which are listed in Table 2. Many of these features collectively contribute to the reflection of light, interpreted by our brains as a pattern. Thus, despite the variation among individual plants due to differences in habitat and developmental stage, it is possible, with practice, to recognize Cryptantha species at a glance (Figure 2A-D). This wholistic, pattern recognition method is how most botanists and hobbyists identify plants. Until one develops a wholistic pattern recognition familiarity with Cryptantha, one must rely instead on analytic methods, especially taxonomic keys and diagnostic descriptions. Interestingly, relatively few of the characters listed in Table 2 are actually used in keys. Among these major features in identification are plant duration, branching pattern and position, bract presence, pedicel length, calyx length (in flower and fruit). Crossosoma 35(1). Spring-Summer 2009 7 Table 2. Some characters used to distinguish between Cryptantha taxa. those in bold most often used in taxonomic keys. calyx/sepal shape, calyx vestiture/trichome type, and corolla limb width (Table 2). How ever, variation in nutlet morphology is most important in Cryptantha identification, including nutlet number (per fruit), size, shape, and sculpturing. (Why nutlet variation is so great in this complex is a question of great interest regarding the evolution of the group.) Thus, it is critical to collect Cryptantha specimens at a stage w hen at least some mature fruits are present. Identification of immature plants can be quite problematic. Observations of most of these key characteristics require the use of a good dissecting microscope and basic dissecting tools. In addition, we have found two procedures to be of value in identification of Cryptantha (and of related taxa, such as Amsinckia , Pectocarya. and Plagiobothrys). First, if nutlets adhere to and are difficult to remove from the gynobase, boiling the fruits for 1-2 minutes will usually loosen them, allow ing them to be removed intact (Figure 2E). Second, to clearly see details of the fruits or calyx, it is valuable to place them on a glass microscope slide covered with a strip of double-stick tape and a label with herbarium voucher information (Figure 2F). Note that in order to get an accurate count of nutlet number, care must be taken both to choose fruits that are not too mature (i.e., before the nutlets or entire fruits have naturally fallen away) and to avoid dislodging nutlets prematurely during dissection. MATERIALS AND METHODS Herbarium specimens or live plants (subsequently vouchered) were photographed with a Nikon Coolpix 4400 or a Sony digital camera. Close-ups of vegetative parts, flowers, or fruits were photographed either using a Wild M3Z dissecting scope with a Nikon Coolpix 995 digital camera or using a Visionary Digital Imaging System (consisting of an FX2 lighting system, Infinity Optics Long Distance Microscope. SolMate trans-illuminator. Precision Linear Camera Controller. 10 megapixel digital camera, and a devoted computer workstation with Helicon Focus software). Appendix Plant duration Stem type Stem vestiture / trichome type Branch number / position Internode length Leaf position Leaf size Leaf shape Leaf vestiture / trichome type Inflorescence position Flower internode length Bract presence / morphology Pedicel length Calyx length (flower vs. fruit) Calyx / Sepal shape Calyx vestiture / trichome ty pe CalvTt posture Calyx orientation Corolla tube length Corolla limb width Corolla color Fomices color Gynobase size / shape Sty le length Nutlet number per fruit Nutlet size Nutlet shape / base / margin / apex Nutlet sculpturing Nutlet symmetry Nutlet ventral groove shape 8 C.rossosoma 35(1 ), Spring-Summer 2009 Figure 2. A-D: Species of Cryptantha, illustrating that many do look different and can be site- identified A: C. maritima B: C. nevadensis. C: C. racemosa. D: C. utahensis E: Boiling of fruits for 1-2 minutes will loosen nutlets. F: Microsope slides with double-stick tape and labels for observing Cryptantha nutlets. Crossosoma 35(1), Spring-Summer 2009 9 1 lists herbarium voucher accessions for all Ciyptantha images (except for a few field photographs), by figure number and letter. Dichotomous keys were modified from pre- existing keys from Munz and Keck (1959). Munz (1974). W.A. Kelley and Wilken (1993). and R.B. Kelley (in prep). For current nomenclature and descriptions, we are especially appreciative of an early version of R.B. Kelley’s treatment of the genus for the upcoming second edition of The Jepson Manual. Information on bioregions was obtained from R.B. Kelley (in prep) and from W.A. Kelley and Wilken (1993). Count} and regional distributions were obtained from herbarium vouchers listed on the Consortium of California Herbaria (2009: see Rosatti 2003) for the region of Southern California as defined in this paper: however, identifications were not confirmed by us, except in a few' cases. Information on sensitive or rare taxa was obtained from the California Native Plant Society Inventory (2009). cited in the text as a CNPS listing, both in descriptions and the Conservation Status and Ecology of Southern California Cryptantha section. The following keys, plus multi-entry (polvthetic) keys, numerous images, distribution maps, and nomenclatural information, are accessible at our Cryptantha web site: http://www.sci.sdsu.edu/plants/cryptantha. Our aim is to provide a service for both professionals and amateurs in the identification of this fascinating group of plants. We provide an initial key to previously recognized groups (variously treated in the past as subgenera or separate genera): Eremocarya, Greene ocharis , Kiynitzkia, and Oreocarya. We do not include detailed descriptions of taxa. but these will be forthcoming in the second edition of The Jepson Manual (R.B. Kelley in prep). We cite only the diagnostic features of each taxon, its general and county distribution, pertinent ecological information, and conservation status, where applicable. Species diagnoses and plates appear in the order presented in keys and not in alphabetical order so as to more clearly illustrate and compare diagnostic features. For ease of use. each major group and individual taxa are numbered in the keys and descriptions. Technical terms that we use include dorsal, for the back side of a nutlet, i.e.. the side facing away from the gvnobase. and ventral, for the inner side of a nutlet, facing the gynobase. (See Ciyptantha web site: Keys for a complete list of terminology and their definitions.) Corolla width is always measured from lobe to lobe of the corolla limb, the expanded portion at the apex of the tube or throat. We do not include distribution maps (due to space concerns), but encourage all to do a specimen search for selected taxa on the Consortium of California Herbaria. 10 Crossosoma 35(1), Spring-Summer 2009 KEYS TO CRYPTANTHA OF SOUTHERN CALIFORNIA The 49 species and 56 taxa (including varieties) of Cryptantha occurring in Southern California are listed in Table 3. The initial key, below, segregates taxa into “groups” that correspond to previously recognized genera or subgenera. The Krynitzkia group is further divided into two keys: one for smooth nutlets and one for rough nutlets. Table 3. The 49 species and 56 taxa of Cryptantha in Southern California. Taxonomy after R. B. Kelley, in prep. *= Eremocarya ; **= Greeneocharis ; ***= Oreocaryw, all others “ Krynitzkia ”. CNPS Inventory listings in bold C. affinis C. gracilis C. nubigena *** C. angustifolia C. hoffniannii *** C. oxygona C. barbigera var. barbigera C. holoptera 4.3 C. pterocarya var. cycloptera C. barbigera var fergusoniae C. humilis *** C. pterocarya var. pterocarya C. cmerea var abortiva *** C. inaecpiata C. pterocaiya var pur pus ii C. circumscissa var circumscissa ** C. intermedia var. intermedia C. racemosa C. clevelandii var. c/evelandii C. leiocarpa C. recurvata C. clevelandii var, fiorosa C. lepida * C. roosiorum *** 1B.2 C. clokeyi 1 B. 1 C. maritima var. maritima C. scoparia 4.3 C. confertiflora *** C. micrantha * C. similis * * C. corol/ata C. micromeres C. simulans C. costata 4.3 C. microstachvs C. sparsiflora C. decipiens C. mohavensis C. torreyana C. dumetorum C. muricata var denticulata C. traskiae 1 B.2 C. echinella C. muricata var jonesii C. tumulosa *** 4.3 C.flaccida C. muricata var. muricata C. utahensis C. flavoculata *** C. nemaclada C. virginensis *** C. gander i 1 B. 1 C. nevadensis var. nevadensis C. watsonii C. glomeriflora 4.3 C. nevadensis var. rigida Key to Cryptantha Groups 1. Plants annual, gen wider than tall, often rounded to cushion-like; taproot red or purple when dry (often staining herbarium paper); flower bracts present 2. Calyx basally fused, tubular, tube circumscissile in fruit Group 1: Greeneocharis , p. 11 2' Calyx of distinct sepals, intact in fruit Group 2: Eremocarya , p. 13 U Plants annual, biennial, or perennial, gen taller than wide, rarely rounded to cushion-like; taproot usually not red or purple when dry'; flower bracts absent or rarely present 3. Plants biennial or perennial; vegetative leaves basal or tufted; nutlets smooth, rugose, or scabrous, apically broadly rounded to obtuse; ventral groove apex well below nutlet apex Group 3: Oreocarya , p. 13, 15 3’ Plants annual or rarely perennial; vegetative leaves cauline; nutlets smooth, papillate, tuberculate, or muricate, apically narrowly acute to acuminate; ventral groove apex almost to nutlet apex Group 4: Krynitzkia , p. 20 Crossosoma 35(1). Spring-Summer 2009 11 Group 1: Greeneocharis \ Piptocalyx \ The two species of Greeneocharis (named after Edward Lee Greene. 1843-1915) are readily distinguished by the combination of red to purple-pigmented roots, bracteate flowers, and a calyx that is basally synsepalous (sepals fused), splitting transversely at fruit maturity, a condition termed circumscissile dehiscence. Interestingly, the only other close relatives with such a calyx dehiscence are two species of Plagiobothrys: P. arizonicus and P. nothofulvus. See Figures 29A.B for comparative nutlet images in the Greeneocharis group. 1. Corolla 1-2 mm wide, fornices tiny, white 1. C. circumscissa var. circumscissa 1’ Corolla 3-6 mm wide, fomices evident, yellow 2. C. similis 1. Cryptantha circumscissa var. circumscissa (Hook. & Am.) I.M. Johnst. (Figure 3 A-E. p. 12 ) Common name: Circumscissile cryptantha. Distribution in Southern California: Southwestern California region (eastern South Coast. Transverse Ranges, and eastern Peninsular Ranges). Desert province. White and Invo mountains: Imperial. Inyo, Kern. Los Angeles, Mono. Riverside, Santa Barbara, San Bernardino. San Diego, and Ventura counties. Conservation Status: None. Diagnosis: Plants (at maturity) generally wider than tall with red-purple roots and bracteate flowers, distinguished from the Eremoccnya group in having circumscissile calyces and from C. similis by the smaller (1-2 mm wide) corolla. Note: C. circumscissa var. rosulata. the ground rose cryptantha (CNPS IB. 2 listing), has a limited distribution in subalpine habitats of the Sierra Nevada. Inyo and Tulare Cos., outside Southern California. 2. Cryptantha similis K. Matthew & PH. Raven (Figure 3F-J, p.12) Common name: Dome cryptantha. Distribution in Southern California: Southwestern California region (northern San Gabriel and western San Bernardino mountains), southwestern Mohave Desert region: Kern, Los Angeles, and San Bernardino counties. Conservation Status: None. Diagnosis: Plants (at maturity) generally wider than tall with red-purple roots and bracteate flowers, distinguished from the Eremocatya group in having circumscissile calyces and from C. circumscissa by the larger (3-6 mm w ide) corolla. [C. circumscissa (Hook. & Arn.) I.M. Johnst. in part, sensu W.A. Kelley and Wilken 1993] These two taxa are distinguished only by corolla size and color of fornices. The range of Cryptantha similis is much more restricted, but appears to overlap, with that of C. circumscissa. 12 Crossosoma 35(1), Spring-Summer 2009 Figure 3. A-E: Crvptantha circumscissa A: Whole plant, showing small height relative to width. B-C: Inflorescence and flowers. Note bracts and small corolla D: Circumscissal calyx. E: Nutlet, dorsal (left), ventral (middle), and lateral (right) views. F-J: C. similis. F: Whole plant. G: Flowering shoot. Note wide (3-6 mm) corolla and floral bracts H: Circumscissal calyx. I: Stem close-up, showing appressed and spreading trichomes. J: Nutlet, dorsal (left) and ventral (right) views. Crossosoma 35( 1 ). Spring-Summer 2009 13 Group 2: Eremocarya The two species of Eremocarya (from eremo< desert + carya , fruit) are distinguished by the combination of red to purple-pigmented roots, bracteate flowers, and an aposepalous calyx (sepals not fused). Thus, it is the latter feature that distinguishes this group from Greeneocliaris. See Figures 29C.D for comparative nutlet images in the Eremocarya group. 1. Corolla limb 1—3.5 mm wide 3. C. lepida I’ Corolla limb 0.5-1. 2 mm wide 4. C. mi era nth a 3. Cryptantha lepida (A. Gray) l.M. Johnst. (Figure 4A-E, p.14) Common name: Elegant cryptantha. Distribution in Southern California: Southwestern California region (Peninsular and Transverse Ranges). Desert province: Imperial. Kern. Los Angeles. Riverside. San Bernardino. San Diego counties. Conservation Status: None. Diagnosis: Plants (at maturity) generally wider than tall with red-purple roots and bracteate flowers, distinguished from the Greeneocharis group in having non-circumscissile calyces and from C. micrantha in having larger (1-3.5 mm wide) corollas. [C. micrantha (Torrev) l.M. Johnst. in part . sensu W.A. Kelley and Wilken 1993] 4. Cryptantha micrantha (Torrev) l.M. Johnst. (Figure 4F-J. p. 1 4) Common name: Purple Root cryptantha. Distribution in Southern California: mostly Desert province: Imperial. Inyo, Los Angeles. Riverside. San Bernardino. San Diego counties. Conservation Status: None. Diagnosis: Plants (at maturity) generally wider than tall with red-purple roots and bracteate flow ers, distinguished from the Greeneocharis group in having non-circumscissile calyces and from C. lepida in having smaller (0. 5-1.2 mm wide) corollas. Note that some overlap occurs in the overall corolla width of these two species. However, within a plant, most flowers fit the range of their respective species. The two taxa also seem to occupy different habits, C. lepida in montane and desert transition regions, occasionally cismontane washes, and C. micrantha in lower elevation desert regions, generally flats and alluvial slopes, often sandy soils, with creosote bush scrub. Group 3: Oreocarya The species of Oreocary;a ( oreo . mountain + carya , fruit) are distinctive in being biennial or perennial taxa with a generally w ell formed and persistent rosette of leaves. Many of these taxa occur at high elevations. In addition, the nutlets in this group tend to be larger than other Cryptantha species, usually with characteristic rounded apices and ventral grooves that do not extend to the nutlet apex. See Figure 26 for comparative nutlet images in the Oreocarya group. 14 Crossosoma 35(1), Spring-Summer 2009 Figure 4. A-E: Cryptantha lepida. A: Whole plant. B: Inflorescence; note floral bracts. C: Flower, showing larger (1-3.5 mm wide) corolla. D: Fruit, with 4 nutlets removed; note unfused sepals (calyx aposepalous) E: Nutlet, dorsal, ventral, and lateral views (left to right) F-J: C. mkrantha. F: Whole plant, wider than tall G: Inflorescence, with floral bracts. H: Flower, showing small (0.5-1 mm wide) corolla. I: Fruit, with unfused sepals. J: Nutlets, two forms: rough (left, dorsal and ventral views) and smooth (right, dorsal and ventral views). Crossosoma 35(1), Spring-Summer 2009 15 Key to Oreocaiya Group 1. Corolla tube > sepals 2. Corolla and fornices yellow; nutlet back smooth, groove closed, ventral groove margins ± overlapping 5. C. confertiflora 2' Corolla white, fornices light yellow; nutlet back rough, groove open, narrowed near middle, ventral groove margins elevated 6. C. flavoculata 1 ’ Corolla tube = or < sepals 3. Stems prostrate to ascending; nutlet dorsal side strongly convexlv curved, surface smooth; groove closed, margins generally overlapping 7. C. cinerea var. abortiva 3' Stems ± erect (generally cespitose in C. humilis ); nutlet back relatively straight, surface smooth or rough; groove closed or open, margins not overlapping 4. Stems 1-2 cm tall. < or = basal leaves; basal leaves generally < 1 cm .... 8. C. roosiorum 4’ Stems > 2 cm tall. > basal leaves; basal leaves generally » 1 cm 5. Nutlet lanceolate to lance-ovate, inner surface (between groove and margin) generally smooth, shiny 9. C. nubigena S’ Nutlet ovate, inner surface generally rough or wrinkled 6. Edge of nutlet groove not elevated; nutlet sculpturing finely rugose-tuberculate on dorsal and ventral surfaces 10. C. humilis 6’ Edge of nutlet groove elevated; nutlet sculpturing roughly rugose 7. Pedicel 0.5-2 mm in fruit; calyx lobes 4.5-8 mm in fruit; trichomes on stem and leaves white 1 1 . C. hoffmannii 7 ’ Pedicel 2-6 mm in fruit; calyx lobes 7-1 1 mm in fruit; trichomes on stem and leaves yellow to brown 8. Biennial or short-lived perennial herbs; caudex not woody; nutlets prominently dorsally ridged or keeled 12. C. virginensis 8' Long-lived perennial herbs; caudex woody; nutlets not prominently keeled, dorsal ridge rounded 13. C. tumulosa 5. Cryptantha confertiflora (Greene) Payson (Figure 5A-B. p.16) Common name: Yellow-flowered cats eye. Distribution in Southern California: Southwestern California region (San Bernardino Mountains), Desert mountains: Inyo. Kern. Mono. San Bernardino counties. Conservation Status: None. Diagnosis: Corolla yellow, tube exceeding calyx in length; nutlets smooth and shiny w ith closed ventral groove with overlapping margins. 6. Cryptantha flavoculata (A. Nelson) Payson (Figure5C-D. p. 16) Common name: Yellow-eyed cat s eye. Distribution in Southern California: Desert mountains; Inyo. Mono. San Bernardino counties. Conservation Status: None. Diagnosis: Corolla white with prominent, bright yellow fornices. tube exceeding calyx in length: nutlets w ith rough sculpturing and open ventral groove w ith elevated margins. 7. Cryptantha cinerea var. abortiva (Greene) Cronquist (Figure 5E-H. p.16) Common name: Bow-nut cat's eye. Distribution in Southern California: San Bernardino. Desert mountains: Inyo, Los Angeles. Mono, San Bernardino counties. Conservation Status: None. Diagnosis: Stems prostrate to ascending: nutlets strongly convex-curved dorsally, smooth and shiny with a closed nutlet groove. [C. jamesii (Torrey) Payson var. abortiva (Greene) Pay son] 16 C.rossosoma 35(1), Spring-Summer 2009 Figure 5. A-B: Cryptantha confertiflora. A: Inflorescence, showing corolla tube longer than calyx B: Nutlet, dorsal (left) and ventral (right) views, showing smooth surface and closed groove. C-D: C. flavoculata. C. Inflorescence, showing long corolla tube D: Nutlet, dorsal (left) and ventral (right ) views, showing rough surface and open groove. E-H: C. cinerea var. abortiva E: Plant, showing prostrate to ascending stem habit. F: Nutlet, side view, showing strongly curved dorsal surface. G: Nutlet, dorsal view H: Nutlet, ventral view. Note: images at A, C, and E courtesy of Gary' Monroe. Crossosoma 35(1), Spring-Summer 2009 17 8. Cryptantha roosiorum Munz (Figure 6 A-B. p. 18) Common name: Bristlecone cat's eye. Distribution in Southern California: Inyo Mountains: Inyo County. Conservation Status: CNPS List IB. 2. Diagnosis: Stems short, often equal in length with the basal leaves and generally no greater than 2 cm tall: basal leaves no greater than 1 cm long. It has a very restricted distribution in high elevation limestone soils of bristle-cone pine and limber pine forests. 9. Cryptantha nubigena (Greene) Pavson (Figure 6C-D. p.18) Common name: Sierra cat's eye. Distribution in Southern California: White and Inyo mountains. Desert mountains: Inyo. Mono. San Bernardino counties. Conservation Status: None. Diagnosis: Stems elongate, much longer than leaves: nutlets lanceolate to lance-ovate with smooth, shiny dorsal and ventral surfaces. 10. Cryptantha hiunilis (Greene) Pavson (Figure 6E-F. p. 18 ) Common name: Low cat's eye. Distribution in Southern California: northern Desert mountains: Inyo. Mono counties. Conservation Status: None. Diagnosis: Stems elongate, much longer than leaves: nutlets ovate, sculpturing finely rugose on dorsal and ventral surfaces, margin of the ventral groove non-elevated. 11. Cryptantha hoffmannii I.M. Johnston (Figure 7A-D. p. 19) Common name: White Mountain cat's eye. Distribution in Southern California: White and Inyo mountains, northern Desert mountains: Inyo, Mono. San Bernardino counties. Conservation Status: None. Diagnosis: Biennial or short- lived perennial: pedicel 0.5-2mm in fruit: calyx 4.5-8 mm long; nutlet coarsely rugose, ventral groove margin elevated. It is distinguished from the closely associated species C. virginensis and C. tumulosa by the snowy-white trichomes and the smaller pedicel and fruiting calyx. It is distinguished from C. humilis by the coarsely rugose sculpturing on the nutlet and elevated margin of the nutlet groove. [C. virginensis (M.E. Jones) Payson in part, sensu W.A. Kelley and Wilken 1993] 12. Cryptantha virginensis (M.E. Jones) Payson (Figure 7E-G. p. 19 ) Common name: Virgin River cat's eye Distribution in Southern California: White and Inyo mountains, northern and eastern Mojave Desert mountains; Inyo. Mono. San Bernardino counties. Conservation Status: None. Diagnosis: Biennial or short lived perennial: caudex slender and short-lived; pedicel 2-5 mm long in fruit; calyx 7-11 mm long; nutlet roughly rugose, dorsally carinate, ventral groove margin elevated. It differs from C. hoffmannii in its tawny, yellow-brown trichomes, the longer pedicel and calyx, and the dorsally carinate (keeled) nutlet. 13. Cryptantha tumulosa (Payson) Payson (Figure 7H-I. p. 19) Common name: New York Mountain cat s eye. Distribution in Southern California: eastern Mojave Desert, northern and eastern Desert mountains: Inyo. Mono. San Bernardino counties. Conservation Status: CNPS List 4.3. Diagnosis: Perennial: caudex woody: pedicel 3-6 mm long in fruit, calyx 7-10 mm long in fruit, nutlet roughly rugose, not carinate, ventral groove margin elevated. It is 18 C.rossosoma 35(1), Spring-Summer 2009 tnutlets landeolate ■lance-ovate; ventralfjJ|| Surface smooth, shinvj&^j Figure 6. A-B: Cryptantha roosioruni A: Whole plant, showing short stature and small leaves. B: Nutlet, dorsal (left) and ventral (right) views C-D: C. nubigena. C: Whole plant D: Nutlet, dorsal (left) and ventral (right) views Note smooth, shiny ventral surface. E-F: C. hunulis. E: Whole plant. F: Nutlet, dorsal (left) and ventral (right) views. Note finely rugose surface, edge of ventral groove not appreciably elevated leaves < 1 cm long stems cm Crossosoma 35(1), Spring-Summer 2009 19 dorsal keel edge of groove elevated f dorsal keel absent or ^ rounded 1 C. virginensis edge of groove elevated j C. tumul os a Figure 7. A-D: Cryptantha hoffmannii. A: Whole plant B: Stem trichomes. white in color C: Calyx in fruit, 4 5-8 mm long. D: Nutlet, dorsal (left) and ventral (right) views. Note elevated ventral groove edge E-G: C. virginensis. E: Whole plant. F: Stem trichomes. yellow to brown in color. G: Nutlet, dorsal (left) and ventral (right) views. Note dorsal keel. H-l: C. tumulosa. H: Whole plant. I: Nutlet, dorsal (left) and ventral (right) views. Note absence of dorsal keel 20 Crossosoma 35(1), Spring-Summer 2009 distinguished from closely associated C. hoffmannii, by its tawny, yellow-brown trichomes, and the longer (in fruit) pedicel and calyx. It differs from C. virginensis in having a prominently woody caudex and nutlets with a rounded dorsal ridge. Group 4: Krynitzkia The species of Kiynitzkia (origin unknown, presumably named as a commemorative), as circumscribed by Johnston (1927) and Payson (1927), are almost certainly an unnatural, paraphyletic group (Hasenstab 2009). Almost all members are annuals (very rarely perennial) with mostly cauline leaves. The nutlets in this group are quite variable in sculpturing, number, and heteromorphism, but all have an acute to narrowly acute apex and a ventral groove that extends almost to the apex. See Figures 27, 28, 29E-H, and 30 for comparative nutlet images in the Kiynitzkia group. For ease of identification, Kiynitzkia is divided into two groups, based on general nutlet sculpturing. Key to Groups of Krynitzkia 1. All nutlets of fruit smooth and shiny Krynitzkia Smooth Nutlet Group, below I’ At least one nutlet of fruit rough: papillate, tuberculate, and/or inuricate Krynitzkia Rough Nutlet Group, p. 29 Kry nitzkia - Smooth Nutlet Group (See Figure 27 for comparative nutlet images) [Note: an asterisk indicates that taxon also appears in the Krynitzkia Rough Nutlet key ] 1. Nutlet asymmetric, ventral groove off-center from nutlet body 2. Nutlet ventral groove strongly off-center, appearing marginal; nutlets 4 per fruit; flowers in elongate inflorescence units 14. C. affinis 2’ Nutlet attachment scar edges slightly off-center, nearer one margin than the other; nutlet 1 per fruit; flowers in globose inflorescence units 15. C. glomeriflora 1 ’ Nutlet symmetric, ventral groove central 3. Some calyx trichomes apically hooked, sometimes basally encrusted with exudate; nutlet 1 per fruit 4. Nutlet lance-ovate with swollen base, rounded margin, and long acuminate (caudate) apex; corolla limb 1-5(6) mm wide; calyx trichomes usually basally encrusted with exudate; common, widespread 16. C. flaccida 4’ Nutlet compressed ovate with dorsally flattened base, sharp angled margin, and ± acute apex; corolla limb 0 5-1(1 .5) mm wide; calyx trichomes not basally encrusted with exudate; uncommon 17. C. sparsifiora 3’ Calyx trichomes ± straight, rarely wavy, either erect, spreading, or reflexed, not apically hooked nor basally encrusted with exudate; nutlets 1—4 per fruit 5. Calyx sepal tips with reflexed bristles; corolla limb 0.5—1 mm wide .... 18. C. nemaclada 5’ Calyx sepal tips with bristles mostly spreading or ascending; corolla limb 0.5-7. 5 mm wide 6. Sepals 6-10 mm in fruit; longest sepal bristles 3^4 mm in fruit 19. C. ganderi 6" Sepals 1-6 mm in fruit; longest sepal bristles < 3 mm in fruit 7. Flowers gen subtended by bracts 8. Stems gen prostrate to decumbent, greenish when in fruit; nutlets 3-4 per fruit (all smooth) 20. C. leiocarpa 8’ Stems gen erect to ascending, reddish brown when in fruit; nutlet 1 per fruit and smooth (or 2 per fruit, 1 smooth, 1 rough) * 21. C. maritima Crossosoma 35(1), Spring-Summer 2009 21 7’ Flower bracts absent 9. Calyx obconic at base; calyx trichomes dense, straight, ascending, relatively soft, sepal midvein hairs slightly longer, but not bristly 22. C. gracilis 9' Calyx rounded at base, with both appressed. soft trichomes and ascending to spreading, stiff, bristle-like trichomes 10. Sepals < 1 mm in fl. 1-2 mm in fruit 23. C. microstachys 10’ Sepals > 1 mm in fl. >3 mm in fruit 11. Nutlets 1-1 .5 mm long, 1^1 per fruit C. ctevelandii 12. Pedicels 0-0.5 mm, becoming erect. ± appressed to stem in fruit; calyx (2.5)3— 3.5 mm in fruit, narrow at base, sepals linear in fruit; corolla limb 1-2.5 mm wide; corolla fornices pale yellow; nutlets 1-2 per fruit 24. C. ctevelandii var. ctevelandii 12’ Pedicels 0.5—1 mm. ± inclined, not appressed to stem in fruit; calyx 3^4(4 5) mm in fruit, ± wide at base, sepals ovate in fruit; corolla limb (2)2. 5-5 mm wide; corolla fornices yellow; nutlets 3^ft 1) per fruit 25. C. ctevelandii var .florosa 1 1 1 Nutlets 1 .5-2 mm long. 4 per fruit 13. Nutlet margin gen rounded throughout, sometimes angled below middle 26. C. torreyana 13’ Nutlet margin angled (obtuse in cross-section), esp near tip 14. Corolla limb (4)5—7 5 mm wide; nutlets ovate to lance-ovate 27. C. mo Itavens is 14’ Corolla limb 2-3 mm wide; nutlets lanceolate to lance-ovate 28. C. watsonii 14. Cryptantha a ffinis (A. Gray) Greene (Figure 8 A. p.22) Common name: Side-grooved cryptantha. Distribution in Southern California: Southwestern California region (Transverse and Peninsular Ranges); Kern. Los Angeles, Mono. San Bernardino. San Diego, and Ventura counties. Conservation Status: None. Diagnosis: Nutlets 4 per fruit with a very eccentric ventral groove appearing to be marginal; inflorescences with straight clusters of flowers. 15. Cryptantha gtomeriflora Greene (Figure 8B.C. p.22) Common name: Trukee cryptantha. Distribution in Southern California: White mountains; Mono County. Conservation Status: CNPS List 4.3. Diagnosis: Nutlet 1 per fruit with a slightly off-center ventral groove; inflorescence with globose clusters of flow ers. 16. Cryptantha jiaccida (Lehmann) Greene (Figure 8D-E. p.22) Common name: Weak-stemmed cryptantha. Distribution in Southern California: Southwestern California region (western Transverse Ranges, eastern San Bernardino mountains. South Coast. Peninsular Ranges). Tehachapi mountains; Inyo. Kern, Los Angeles, Riverside. Santa Barbara. San Diego. Ventura counties. Conservation Status: none. Diagnosis: Calyx apicallv hooked, trichomes usually basally encrusted with exudate: nutlet 1 per fruit, lance-ovoid with a swollen base, rounded margin, and long acuminate apex. 22 Crossosoma 35( 1 ), Spring-Summer 2009 Figure 8. A: Cryptantha affinis , nutlet, ventral view, showing strongly off-center ventral groove. B-C: C. glomeriflora. B: Nutlet, ventral view, showing slightly off-center (asymmetric) ventral groove. C: Inflorescence unit, a tightly clustered, globose circinate cyme D-E: C. flaccida. D: Sepal, showing hooked (uncinate) trichomes (arrow) E: Nutlet, dorsal, ventral, and lateral views (left to right). Note lanceolate shape, globose base, rounded margin, and elongate, acuminate (caudate) apex. F-G: C. sparsifloru. F: Sepal, showing hooked (uncinate) trichomes (arrow). G: Nutlet, dorsal, ventral, and lateral views (left to right). Note ovate shape, dorsalty flattened base, sharp-angled margin, and short, acute apex Crossosoma 35(1). Spring-Summer 2009 23 17. Cryptantha sparsiflora (Greene) Greene (Figure 8F-G. p.22) Common name: Few-flowered cryptantha. Distribution in Southern California: Southwestern California region (South Coast). Tehachapi mountains; Kern. Ventura counties. Conservation Status: none. Diagnosis: Calyx trichomes apically hooked, not basally encrusted w ith exudate; nutlet 1 per fruit, ovate with dorsally flattened base, sharp-angled margin, and acute apex. [C. flaccida ( Lehmann) Greene in part. sensu W.A. Kelley and Wilken 1993] 18. Cryptantha nemaclada Greene (Figure 9A-B. p.24) Common name: Colusa cryptantha. Distribution in Southern California: Southwestern California region (w estern Transverse Ranges). Tehachapi mountains: Kern. Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Sepal apices with reflexed trichomes; nutlets 1-2 (4) per fruit. 19. Cryptantha ganderi l.M. Johnst. (Figure 9C-D. p.24) Common name: Gander's cryptantha. Distribution in Southern California: western Sonoran Desert; San Diego County (Borrego Valley). Conservation Status: CNPS List 1B.1. Diagnosis: Sepals relatively long (6-10 mm) with long (3-4 mm) sepal bristles: nutlets large (2-2.5 mm long). 20. Cryptantha leiocarpa (Fisher & C. A. Mever) Greene (Figure 9E. 9G. p.24) Common name: Beach cryptantha. Distribution in Southern California: Southwestern California region (northern South Coast); Los Angeles, Orange. Santa Barbara, and Ventura counties. Conservation Status: none. Diagnosis: Plant decumbent; stems greenish at maturity: flowers bracteate: nutlet 1 per fruit. 21. Cryptantha maritima (Greene) Greene var. maritima (Figure 9F. 9H. 19A-C. p.24,40) Common name: Maritime cryptantha. Distribution in Southern California: Southwestern California region (South Coast, Channel Islands, Transverse Ranges. Peninsular Ranges). Desert prov ince: Imperial. Inyo. Los Angeles. Riv erside. Santa Barbara. San Bernardino. San Diego, and Ventura counties. Conservation Status: none. Diagnosis: Plants erect; stems reddish at maturity : flowers bracteate: nutlets either 1 per fruit and smooth or 2 per fruit w ith one smooth and one tuberculate. [C. maritima var. pilosa l.M. Johnston is purported to differ from C. maritima var. m. in having dense tufts of soft (pilose) trichomes arising from the calyx, but this feature forms a grade is not diagnostic, sensu R.A. Kelly (in prep): C. maritima var. cedrocensis (Greene) l.M. Johnston, endemic to Cedros Island, Baja California. Mexico, differs in having 4 nutlets (Wiggins 1980)] 22. Cryptantha gracilis Osterh. (Figure 10A-B. p.26) Common name: Slender cryptantha. Distribution in Southern California: Desert province (eastern Mojave Desert and mountains). White and Inyo mountains: Inyo, Mono. San Bernardino counties. Conservation Status: none. Diagnosis: Calyx obconical, with ascending, soft trichomes: nutlet 1 (2.3) per fruit. 3-sided. 24 Crossosoma 35( I ), Spring-Summer 2009 |n lowers! [practeTte] flowers bracteate Jp d Wfganueri [S4mqnUinm ] maritime i Figure 9. A-B: Cryptantha nemaclada A: Calyx, showing short, reflexed bristles near apex B: Nutlet, ventral (left) and dorsal (right) views C-D: C. ganderi. C: Calyx. Note long length D: Nutlet, dorsal view E&G: C. leiocarpa. E: Inflorescence branch, showing bracteate flowers G: Plant from field, showing decumbent branching. Inset: Nutlet, dorsal view F&H: C. maritima F: Inflorescence, showing bracteate flowers H: Plant, showing erect to ascending branches Inset: Nutlet, dorsal view. Crossoso/na 35(1), Spring-Summer 2009 25 23. Cryplantha microstachys (A. Gray) Greene (Figure 10C-E, p.26) Common name: Tejon cryptantha. Distribution in Southern California: Southwestern California region (Transverse Ranges, Peninsular Ranges, South Coast, Santa Catalina Island); Kern. Los Angeles. Orange, Riverside. Santa Barbara, San Bernardino. San Diego, and Ventura counties. Conservation Status: none. Diagnosis: Calyx small, short (< 1 mm in fl. 1-2 mm in fr); nutlet usually solitary. 24. Cryptantha Cleveland" Greene var. Cleveland ii (Figure 10F-H, p.26) Common name: Cleveland's cryptantha. Distribution in Southern California: Southwestern California region (South Coast); Los Angeles, Santa Barbara. San Diego counties. Conservation Status: none. Diagnosis: Flowers ebracteate; pedicels absent to short (0-0.5 mm), becoming appressed to the stem in fruit; calyx basally narrowed with straight, non-reflexed. soft and bristle-like trichomes, the calyx (2. 5)3-3. 5 mm in fruit; corolla 1-2.5 mm wide; nutlets 1-2 per fruit (1-1.5 mm long). 25. Cryptantha Cleveland" Greene var. florosa I.M. Johnst. (Figure 11A-D. p.27) Common name: Coastal cryptantha. Distribution in Southern California: Southwestern California region (South Coast, Channel Islands); Kern, Los Angeles, Orange. Riverside, Santa Barbara. San Bernardino. San Diego, and Ventura counties. Conservation Status: none. Diagnosis: This variety differs in having longer (0.5-1 mm) pedicels that become inclined (not appressed) to the stem in fruit, a longer (3-4 mm long), basally wide calyx (in fruit), a wider (2.5-5 mm) corolla limb, and nutlets 3-4( 1 ) per fruit. [C. Cleveland ii Greene in part . sensu W.A. Kelley and Wilken 1993] 26. Cryptantha torreyana (A. Gray) Greene (Figure 1 1E-J. p.27) Common name: Torrey’s cryptantha. Distribution in Southern California: Southwestern California region (western Transverse Ranges). White mountains; Inyo, Kern, and Ventura counties. Conservation Status: none. Diagnosis: Flowers ebracteate: calyx with a rounded base and straight, non-reflexed trichomes; nutlets symmetrical (1.5-2 mm long). 4 per fruit, w ith rounded margins. 27. Cryplantha mohavensis (Greene) Greene (Figure 12A-B. p.28) Common name: Mohave cryptantha. Distribution in Southern California: Tehachapi mountains, western Mohave Desert, edge of desert only, mostly found along the desert slope of mountains; Inyo, Kern, Los Angeles, San Bernardino counties. Conservation Status: none. Diagnosis: Flowers ebracteate: calyx with a rounded base (1-6 mm long), trichomes straight, non-reflexed; corolla limb relatively large (4-7.5 mm wide); nutlets symmetrical, ovate to lance-ovate (ca. 2 mm long). 4 per fruit, with distinctly angled margins. 26 Crossosoma 35( 1 ), Spring-Summer 2009 \^nw^ostacn^ micros , 'WWle\VlanaiiM ■yy^cleyelanai i wWKi^mandii var. cleyelandii a A 4* A 1 5 , C. gracilis sepals 1-2 mm in fr < 1mm m II corolla fruits appressed / fleam 2.5 mm C. microstachys f • nutlets m Pr 1-1.5 mm Ions JK k C. clevelandii var. clevelandii Figure 10. A-B: Cryptantha gracilis A: Calyx, showing characteristic obconic base and long, soft trichomes. B: Nutlet, dorsal, ventral, and lateral views (left to right). C-E: C. microstachys. C: Flower. Note short calyx. D: Fruit, also with short calyx. E: Nutlet, dorsal (left) and ventral (right) views. F-H: C. clevelandii var. clevelandii. F: Flower, showing small corolla. G: Fruit. Note appressed orientation. H: Nutlet, dorsal (left) and ventral (right) views. Crossosoma 35( 1 ). Spring-Summer 2009 27 Figure 11. A-D: Cryptantha clevelandii var .florosa A: Fruit, showing inclined orientation. B: Corolla, relatively wide in size C: Fruit, showing 4 nutlets. D: Nutlet, dorsal (left) and ventral (right) views. E-J: C. torreyana. E: Whole plants from herbarium specimen. F: Close-up of leaf (left) and stem (right) G: Young inflorescence H: Flower I: Fruit J: Nutlet, dorsal, ventral, lateral, and cross-sectional views (left to right). Note triangular shape and rounded margins. 28 C.rossosoma 35(1), Spring-Summer 2009 ^^orolla 4 - 7 5 mm .wide] [C. mohaveiisis} nutlets lance-ovate to ovate margin sharp-angled C. mohavensis margin sharp-angled I I Jl corolla 1-2 mm wide ◄ ► Figure 12. A-B: Cryptantha mohavensis. A: Nutlet, dorsal, ventral, lateral, and cross-sectional views (left to right). Note ovate shape and rounded ventral face (far right) and sharp-angled margins B: Corolla, with wide limb. C-D: C. watsonii. C: Corolla, with short limb. D: Nutlet, dorsal, ventral, lateral, and cross-sectional views (left to right). Note lanceolate shape and sharp-angled margins. Crossosoma 35(1), Spring-Summer 2009 29 28. Cryptantha watsonii (A. Gray) Greene (Figure 12C-D, p.28) Common name: Watson's cryptantha. Distribution in Southern California: White and Inyo mountains; Inyo, Mono counties. Conservation Status: none. Diagnosis: Flowers ebracteate: calyx with a rounded base (1-6 mm long), trichomes straight, non-reflexed; corolla relatively small (limb 1-2 mm wide); nutlets lanceolate to lance-ovate (1.5-2 mm long). 4 per fruit, with distinctly angled margins. Kn nitzkia - Rough Nutlet Group (See Figure 28. 29E-H, 30 for nutlet images) [Note: an asterisk indicates that taxon appears in the Krynitzkia Smooth Nutlet key or in more than 1 couplet ] 1. One or more nutlets of fruit with margin sharp-angled and flattened as a narrow rim or a narrow to broad w ing 2. Pedicels 1-4 mm long in fruit; plant annual or perennial 3. Nutlets homomorphic, dorsally tuberculate-papillate. margin with narrow, flattened rim or wing; plant usu. annual, w ith major central axis; inflorescence in fruit dense 29. C. holoptera 3’ Nutlets heteromorphic. 1 longer and dorsally papillate to papillate-tuberculate, 2-3 shorter and dorsally papillate-tuberculate. margins with very narrow, flattened rim; plant perennial (but may flower first year), without major, central axis; inflorescence in fruit open 30. C. racemosa 2’ Pedicels 0-1 mm long in fruit; plant annual 4. Nutlet 1 (rarely 2) per fruit; fruit sepals ovate-elliptic, nutlet margin sharp-angled to rimmed, dorsal tubercles minutely spinulose 31. C. utahensis 4’ Nutlets usu. 4 per fruit; fruit sepals linear-lanceolate, nutlet margin winged along entire margin, dorsal tubercles, if present, not spinulose 5. At least some nutlets with prominent, strongly lobed wings C. pterocarya 6. Nutlets heteromorphic, 3 nutlets winged, 1 not winged 7. Wing of winged nutlets very broad 32. C. p. var. pterocarya 7’ Wing of w inged nutlets narrow, knifelike 33. C. p. var .purpusii 6’ Nutlets homomorphic, all 4 nutlets with prominent wings 34. C. p. var. cvcloptera 5’ Nutlets sharp-angled, rimmed, or narrowly winged (not strongly lobed) 8. Nutlets homomorphic 9. Corolla limb 4—9 mm wide; calyx in fruit 3-4 mm long; nutlets 2-2.5 mm long, narrowly winged (wings sometimes toothed), dorsal surface coarsely tuberculate; flowers ebracteate 35. C. oxygon a 9’ Corolla limb 1-3 mm wide; calyx in fruit 4-6 mm long; nutlets 1.5—2 mm long, narrowly winged (wings sharp, never toothed), dorsal surface finely tuberculate; flowers bracteate (bracts deciduous) 36. C. costata 8’ Nutlets heteromorphic by size, one distinctly larger than the other three 10. Nutlet margin rimmed and knife-like to narrowly winged; small nutlets ca 1.3 mm long 37. C. inaequata 10’ Nutlet margin sharp-angled to slightly rimmed (usually rounded); small nutlets ca. 1 mm long * 38. C. angustifolia 1’ All nutlets of fruit w ith margin rounded, ridged, or sharp-angled (acute to obtuse), but not flattened as a rim or wing 11. Nutlets 2-4 per fruit, heteromorphic by size, one nutlet larger than the other(s) 12. Nutlets 4 per fruit, similar in sculpturing 13. Plants erect; nutlets lance-ovate, small nutlets ca. 1 mm long, finely papillate, ventral groove narrow * 38. C. angustifolia 30 Crossosoma 35(1 ), Spring-Summer 2009 13’ Plants sprawling; nutlets narrowly lanceolate, small nutlets ca. 2 mm long, tuberculate-papillate, ventral groove wide 39. C. dumetorum 12’ Nutlets 2-4 per fruit, different in sculpturing, the larger nutlet smooth to mostly smooth, the smaller nutlet(s) papillate to tuberculate 14. Nutlets 4 per fr, < 1 mm long, triangular-ovate; flower bracts absent; calyx in fruit sub-globose with hooked trichomes, sepals 1—1.5 mm long 40. C. micromeres 14’ Nutlets 2 per fruit, 1.5-2 mm long, oblong-lanceolate; flower bracts present, calyx in fruit oblong with straight trichomes, sepals 2-3 mm long * 21. C. maritima 11’ Nutlets 1-4 per fruit, homomorphic in size and sculpturing 15. Nutlets lanceolate (length:width ratio >3:1) 16. Nutlet 1 (-2) per fruit 17. Mature fruit strongly recurved from stem; nutlet incurved 41. C. recurvata 17’ Mature fruit not recurved from stem; nutlet(s) not incurved 18. Corolla limb 1-1.5 mm wide, deserts 42. C. decipiens 18’ Corolla limb 3-5 mm wide, cismontane 43. C. eorollata 16’ Nutlets usu. 4 per fruit 19. Calyx 6-10 mm long; nutlets coarsely tuberculate C. nevadensis 20. Plants 20-60 cm tall; secondary branches sprawling; sepals strongly recurved at tips; corolla limb 1-2 mm wide 44. C. nevadensis var. nevadensis 20’ Plants <20 cm tall; secondary branches ascending; sepals erect to slightly recurved at tips; corolla limb 2^1 (5) mm wide 45. C. nevadensis var. rigida 19’ Calyx 4-6 mm long; nutlets finely tuberculate-spinulose 46. C. scoparia 15’ Nutlets lance-ovate, ovate, or triangular-ovate (length: width ratio < 3: 1 ) 21. Nutlet dorsal surface with at least a faint median ridge C. muricata 22. Corolla limb 1—2.5 mm wide: nutlets brownish 23. Stem bearing, from top to below middle, many short, flower-bearing branchlets from a larger, primary axis, forming an elongate, leafy, thyrse inflorescence 47. C. muricata var. jonesii 23’ Stem loosely and sparsely branched, without a larger primary axis 48. C. muricata var. denticulata 22' Corolla limb 3-8 mm wide, nutlets grayish 49. C. muricata var. muricata 21’ Nutlet dorsal surface lacking median ridge 24. Corolla limb 3-9 mm wide 25. Sepals 2-3 mm in flower. 3.5-5 mm in fruit 50. C. intermedia var. intermedia 25' Sepals 4—6 mm in flower, 5-10 mm in fruit 51. C. barbigera var. fergusoniae 24’ Corolla limb 1-2 mm wide 26. Nutlets 1—1.5 (1.8) mm long; flowers bracteate; southern Channel Islands 52. C. traskiae 26’ Nutlets 1.5-3 mm long; flowers ebracteate; mainland 27. Nutlet dorsal face flattened to low-convex, densely papillate, sparsely tuberculate 53. C. sinuilans 27’ Nutlet dorsal face convex, densely tuberculate, papillate or not 28. Nutlets ca. 3 mm long, turbercle tips translucent 54. C. clokeyi 28’ Nutlets l .5-2 mm long, turbercle tips not translucent 29. Calyx in fruit 5-6 mm long; nutlets gen 4, ovate, sculpturing finely tuberculate and papillate; corolla limb 1-1.5 mm wide 55. C. echinella 29’ Calyx in fruit 5-10 mm long; nutlets 1-4, lanceolate to ovate, sculpturing coarsely tuberculate to muricate. not papillate; corolla limb 1-2.5 mm wide 56. C. barbigera var. barbigera Crossosoma 35( I ), Spring-Summer 2009 31 29. Cryptantha holoptera (A. Gray) J.F. Macbr. (Figure 13A-C, p.32) Common name: Winged cryptantha. Distribution in Southern California: eastern Mohave Desert, Sonoran Desert: Imperial, Inyo, Los Angeles, Riverside, San Bernardino, and San Diego counties. Conservation Status: CNPS List 4.3. Diagnosis: Annual duration (sometimes perennial); stems with a major axis (inflorescence "dense”); pedicels long (1-4 mm); nutlets homomorphic, tuberculate- papillate. narrowly winged (4 per fruit). 30. Cryptantha racemosa (S. Watson) Greene (Figure 13D-G, p.32) Common name: Bushy cryptantha. Distribution in Southern California: Southwestern California region (northern San Bernardino mountains, eastern Peninsular Ranges), Desert province, northern White and Inyo mountains; Imperial. Inyo. Los Angeles. Mono. Riverside, San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Perennial duration; stems without a single, major axis (inflorescence "open”); pedicels long (1-4 mm): nutlets heteromorphic. papillate to tuberculate-papillate, narrowly winged (4 per fruit). 31. Cryptantha utahensis (A. Gray) Greene (Figure 14A-D. p.33) Common name: Utah/scented cryptantha. Distribution in Southern California: Desert province, White and Inyo mountains; Inyo, Kern, Los Angeles, Riverside. San Bernardino. San Diego counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlet 1 per fruit, with distinctive tuberculate-spinulose processes and a sharp-angled to rimmed margin. 32. Cryptantha pterocarya (Torrey) Greene var. pterocarya (Figure 14E-G, 15A-B. p.33,34) Common name: Fringe nut cryptantha. Distribution in Southern California: Southwestern California region (eastern Peninsular Ranges), Desert province. White and Inyo mountains, often found in rocky uplands; Imperial, Inyo, Kern, Los Angeles, Mono. Riverside, San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, at least some of which have distinctive, strongly lobed wings. This variety is distinguished from the other two by the heteromorphic nutlets, three of which have broad (>0.5 mm), strongly lobed wings and one of which is unwinged. 33. Cryptantha pterocarya (Torrey) Greene var. purpusii Jeps. (Figure 15D-E, p.34) Common name: Argus Mountains cryptantha. Distribution in Southern California: Southwestern California region (Peninsular Ranges, Transverse Ranges). Desert province. White and Invo mountains; Imperial, Inyo, Kern. Los Angeles, Riverside. San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm): nutlets 4 per fruit, at least some of which have distinctive, strongly lobed wings. This variety is distinguished from the other two in having heteromorphic nutlets, three of which have narrow (<0.5 mm), lobed wings and one of which is unwinged. [C. pterocarya (Torrey) Greene in part , sensu W.A. Kelley and Wilken 1993] 32 Crossosoma 35(1). Spring-Summer 2009 Figure 13. A-C: Cryptantha holoptera. A: Fruit, showing relatively long pedicel B: Nutlet, dorsal (left) and ventral (right) views. Note ovate-triangular shape, narrow winge and scattered tubercles. C: Whole plants, showing single, major axis D-G: C. racemosa. D: Whole plants, showing dense banching with no major primary axis E: Fruit, showing long pedicel F: Inflorescence G: Nutlets from a single fruit, dorsal view. Note size heteromorphism. Crossosoma 35( 1 ), Spring-Summer 2009 33 margin sharp-angled ' to slightly winged pterocary'a pterocarya Figure 14. A-D: Cryptantha utahensis. A: Mature, elongate inflorescence B: Inflorescence unit Note narrowly elliptic sepals C: Fruit, showing single nutlet D: Nutlet, dorsal, ventral, lateral, and cross-sectional views (left to right). Note sharp-angled ridge and spinulose sculpturing. E-G: C. pterocarya var. pterocarya. E: Inflorescence, showing swollen, ovoid calyx and small corollas. F: Mature fruit, with four nutlets. G: Close-up of stem (below and right) and leaf (upper left). 34 Crossosoma 35(1), Spring-Summer 2009 C. pterocarya var. purpusii nutlets homomorphic C. pterocarya var. cyl opt era Figure 15. A-B: Cryptantha pterocarya var. pterocarya A: Heteromorphic nutlets of same fruit Left: one of three winged nutlets (dorsal and ventral views). Right: Single unwinged nutlet, dorsal and ventral views. B: Nutlets of a common fruit, heteromorphic: three winged, one unwinged C: C. pterocarya var. cycloptera. Nutlets homomorphic, all winged D-E: C. pterocarya var. purpusii D: Nutlet, dorsal view. Note narrow, lobed wing. E: Nutlets of a common fruit, heteromorphic: three winged, one unwinged. Crossosoma 35( 1 ), Spring-Summer 2009 35 34. Cryptantha pterocarya (Torrey) Greene var. cycloptera (Greene) J.F. Macbr. (Figure 15C, p.34) Common name: Tucson cryptantha. Distribution in Southern California: Southwestern California region (eastern Peninsular Ranges), Desert province. White and Inyo mountains, lower elevations of the desert: Imperial. Inyo. Riverside, San Bernardino. San Diego counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, at least some of which have distinctive, strongly lobed wings. This variety is distinguished from the other two in having homomorphic nutlets, all of which have broad (>0.5 mm), strongly lobed wings. [C. pterocarya (Torrey) Greene in part , sensu W.A. Kelley and Wilken 1993] 35. Cryptantha oxygona (A. Gray) Greene (Figure 16A-E, p.36) Common name: Sharp nut cryptantha. Distribution in Southern California: Southwestern California region (Transverse and northern Peninsular Ranges), western Mohave Desert, Tehachapi mountains. White and Inyo mountains; Inyo, Kern. Los Angeles, Mono. Riverside, San Bernardino. Santa Barbara, and Ventura counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); corolla large (4-9 mm wide); nutlets 4 per fruit (2-2.5 mm long), homomorphic, and narrowly winged (wings sometimes toothed but not strongly lobed). 36. Cryptantha costata Brandegee (Figure 16F-K, p.36) Common name: Ribbed/ashen cryptantha. Distribution in Southern California: Desert province. White and Inyo mountains, in sand dunes; Imperial, Inyo, Riverside. San Bernardino, San Diego counties. Conservation Status: CNPS List 4.3. Diagnosis: Pedicels short (<1 mm); corolla small (1-3 mm wide); nutlets 4 per fruit ( 1 .5-2 mm long), homomorphic, narrowly winged (wings sharp, never toothed or lobed). 37. Cryptantha inaequata I.M. Johnst. (Figure 17A-F. p.37) Common name: Panamint cryptantha. Distribution in Southern California: Mohave Desert, especially eastern; Imperial. Inyo, Riverside. San Bernardino counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, tuberculate-papillate, rimmed to narrowly winged (wings sharp, never toothed or lobed), heteromorphic by size. 1 slightly larger than the other 3. 38. Cryptantha angustifolia (Torrey) Greene (Figure 17G-K, p.37) Common name: Narrow-leaved cryptantha. Distribution in Southern California: Desert province; Imperial, Inyo. Riverside. San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, tuberculate-papillate. margin usually rounded (rarely rimmed to narrowly winged, wings sharp, never toothed or lobed), usually heteromorphic by size, 1 slightly larger than the other 3 (rarely homomorphic). In addition to variation in nutlet morphology in this species, two forms have been noted that vary in corolla limb width and trichome density (personal communication). 36 Crossosoma 35(1), Spring-Summer 2009 Figure 16. A-E: Cryptantha oxygona, A: Whole plant from specimen B: Flower, showing relatively wide corolia C: Fruit D: Fruit with 4, homomorphic nutlets removed. E: Nutlet, dorsal, ventral, and side views (left to right). Note narrow, toothed wing. F-K: Cryptantha costata. F: Whole plant from specimen. G: Plant in field Note large, lanceolate inflorescence bracts. H: Close-up of leaf (left) and stem (right). I: Fruit J: Fruit with sepals removed, showing four nutlets. K: Nutlet, dorsal (left) and ventral (right) views. Crossosoma 35( 1 ), Spring-Summer 2009 37 r. angustifolia X^angustifplui large nutlet small nutlet margin narrowly winged C. inaequata C. inaequata small nutlet nutlets 4 per fruit, heteromorphic C. angustifolia C. angustifolia Figure 17. A-F: Cryptantha inaequata. A: Whole plants from specimen B: Close-up of stem (left) and leaf (right). C: Flower, showing wide corolla limb D: Heteromorphic nutlets, one larger. E: Large nutlet, dorsal, ventral, and side views (left to right). F: Small nutlet, dorsal, ventral, and side views (left to right) Note ovate-triangular shape and narrow wing G-K: C. angustifolia. G: Whole plant in field H: Inflorescence close-up I: Close-up of stem (left) and leaf (right). J: Nutlets of one fruit, showing size heteromorphism Note larger nutlet adherent to gynobase. K: Small nutlet, dorsal, ventral, and side views (left to right). 38 Cross os oma 35(1 ), Spring-Summer 2009 39. Cryptanlha dumetorum (A. Gray) Greene (Figure 18A-G. p.39) Common name: Bush-loving cryptantha. Distribution in Southern California: Desert province. Southwestern California region (northern Transverse Ranges): Inyo, Kern, Los Angeles. Riverside, San Bernardino counties. Conservation Status: none. Diagnosis: Stem habit sprawling, usually climbing in shrubs, foliage very green; nutlets long (2-2.5 mm), lanceolate to lance-ovate, turberculate- papillate, heteromorphic by size ( 1 larger, 3 smaller) with a wide ventral groove. 40. Cryptantha micromeres (A. Gray) Greene (Figure 18H-M, p.39) Common name: Minute-flowered cryptantha. Distribution in Southern California: Southwestern California region (South Coast, Channel Islands. Peninsular Ranges, and Transverse Ranges); Los Angeles, Riverside, San Bernardino, San Diego, Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: Calyx sub-globose with hooked trichomes, 1-1.5 mm long; nutlets small (<1 mm long), heteromorphic. 1 large and smooth to mostly smooth. 3 smaller and tuberculate-papillate. 41. Cryptantha recurvata Coville (Figure 19D-E, p.40) Common name: Bent-nut cryptantha. Distribution in Southern California: Desert province (Mohave, rare in Sonoran), White and Inyo mountains; Inyo, Mono, San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Nutlet 1 (-2) per fruit, strongly recurved from stem, recurved in fruit at maturity, round-margined, lanceolate, and tuberculate-papillate. 42. Cryptantha decipiens (M.E. Jones) A. A. Heller (Figure 19F-H, p.40) Common name: Gravel cryptantha. Distribution in Southern California: Desert province. White and Inyo mountains: Imperial. Inyo. Kern, Los Angeles, Riverside. San Bernardino, San Diego, Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Corolla small (1-1.5 mm): nutlet 1 (-2) per fruit, not recurved from stem at maturity, straight, round-margined, lanceolate, and tuberculate-papillate. 43. Cryptantha corollata (I.M. Johnst.) I.M. Johnst. (Figure 20A-B, p.41) Common name: Coast range cryptantha. Distribution in Southern California: Southwestern California region (Transverse Ranges, northern Peninsular Ranges), Tehachapi mountains; Los Angeles. Riverside. San Bernardino, Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Corolla relatively large (3-5 mm); nutlet 1 (-2) per fruit, not recurved from stem at maturity, straight, round- margined. lanceolate, and tuberculate-papillate. [C. decipiens (M.E. Jones) A.A. Heller in part . sensu W.A. Kelley and Wilken 1993] 44. Cryptantha nevadensis A. Nels. & P.B. Kennedy var. nevadensis (Figure 20C-H, p.41) Common name: Nevada cryptantha. Distribution in Southern California: Desert province. Tehachapi mountains. Southwestern California region (Transverse and Peninsular Ranges). White and Inyo mountains; Imperial. Inyo, Kern, Los Angeles. Mono, Riverside, San Bernardino, San Diego, Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: Secondary branches sprawling, tending Crossosoma 35( 1 ), Spring-Summer 2009 39 Figure 18. A-G: Cryptantha dumetorum. A: Apical shoot of plant, from field Stem is sprawling. B: Stem and leaf close-up. C-D: Inflorescences. Note appressed flowers and fruits. E: Large nutlet, dorsal (left) and ventral (right) views. Note large, open ventral groove F: Small nutlet, dorsal (left) and ventral (right) views. G: Two of four nutlets (small at left, large at right) still attached to gynobase. H-M: C. micromeres H: Whole plant, from specimen. I: Inflorescence. J: Fruit, showing ovoid calyx and hooked trichomes. K: Fruit w ith four nutlets removed. Note heteromorphism. L: Large, mostly smooth nutlet, dorsal (left) and ventral (right) views. M: Small, turberculate nutlet, dorsal, ventral, and side views. 40 Crossosoma 35(1), Spring-Summer 2009 Wjnaniti'na C. recun^ata G miens ciecipiensi mature nutlet strongly incurved C. recurvata ^ mature nutlet W not incurved L l|P C. decipiens lihurj ra]hT?rtrgnfa fiSsttiiW-dl 4 * 0 Figure 19. A-C: Cryptantha maritima. A: Inflorescence, showing bracteate flowers B: Close- up of flowers and fruits. Note dense, straight trichomes. C: Heteromorphic nutlets, dorsal view, smooth (left) and apically rough (right) D-E: C. recurvata. D: Fruit, showing recurved calyx. E: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. Note recurved apex. F-H: C. decipiens. F: Fruit, asceding to inclined (not recurved). G: Flower, showing small corolla. H: Nutlet, dorsal (left) and ventral (right) views. Crossosoma 35( 1 ), Spring-Summer 2009 41 Figure 20. A-B: Cryptanlha corollata. A: Flower, showing large corolla B: Nutlet close- up, dorsal, ventral, and lateral views. Note relatively straight apex C-H: C. nevadensis var. nevadensis. C: Herbarium specimen, showing sprawling stem habit. D: Inflorescence, showing strongly recurved sepal tips E: Close-up of stem (left) and leaf (lower right) F: Calyx of fruit. Note long length G: Flower, showing small (1-2 mm w ide) corolla limb. H: Nutlet close-up, dorsal (left) and ventral (right) views. Note coarse tuberculation. 42 Crossosoma 35(1), Spring-Summer 2009 to crawl through shrubs; calyx relatively long (6-10 mm) with strongly recurved tips; corolla small (1-2 mm wide); nutlets round-margined, lanceolate, 4 per fruit, straight, coarsely tuberculate. 45. Cryptantha nevadensis A. Nels. & P.B. Kennedy var. rigida l.M. Johnst. (Figure 21 A-E, p.43) Common name: Rigid cryptantha. Distribution in Southern California: Tehachapi mountains, Mohave Desert, Southwestern California region (Transverse and Peninsular Ranges), brushy slopes along the edge of the desert; Inyo, Kern, Los Angeles. San Bernardino, Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: Plant +/- erect and self supporting, with ascending secondary branches; calyx with erect to slightly recurved tips; corolla limb larger (2-4 mm wide). [C. nevadensis A. Nels. & P.B. Kennedy in part , sensu W.A. Kelley and Wilken 1993] 46. Cryptantha scoparia Nelson (Figure 2 1F-H, p.43) Common name: Great Basin cryptantha. Distribution in Southern California: White and Inyo mountains: Inyo, Mono counties. Conservation Status: CNPS List 4.3. Diagnosis: Calyx smaller (4-6 mm) and round-margined; nutlets lanceolate. 4 per fruit, straight, finely tuberculate-spinulose. [C. nevadensis A. Nels. & P.B. Kennedy in part , sensu W.A. Kelley and Wilken 1993] 47. Cryptantha muricata (Hook. & Am.) A. Nels. & J.F. Macbr. var . jonesii (A. Gray) l.M. Johnst. (Figure 22A, 22D-F, p.44) Common name: Jones’s cryptantha. Distribution in Southern California: Southwestern California region (South Coast, Transverse and Peninsular Ranges); Los Angeles. Orange, Riverside, San Bernardino, San Diego, Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Corolla smaller (1-2.5 mm); nutlets brownish; stems with a relatively large primary axis bearing many short, flower-bearing branchlets, forming an elongate, leafy, thyrse inflorescence. [C. muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. in part, sensu W.A. Kelley and Wilken 1993] 48. Cryptantha muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. var. denticulata (Greene) l.M. Johnst. (Figure 22B. p.44) Common name: Prickly-nut cryptantha. Distribution in Southern California: Southwestern California region (Transverse and Peninsular Ranges); Los Angeles, Riverside, San Bernardino, San Diego counties. Conservation Status: none. Diagnosis: Nutlets ovate to triangular-ovate, muricate, with at least a faint median ridge on the dorsal surface. This variety is distinguished from the others by the smaller (1-2.5 mm) corolla, brownish nutlets, and stems that are loosely and sparsely branched, without a large, primary axis. [C. muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. in part . sensu W.A. Kelley and Wilken 1993] Crossosoma 35( 1 ), Spring-Summer 2009 43 Figure 21. A-E: Cryptantha nevadensis var. rigidiL A: Herbarium specimen B: Flower close- up. showing corolla (2-4 mm wide) C: Inflorescence with fruits. Note long calyx with slightly- recurved sepal tips D: Fruit, with 4 nutlets removed E: Nutlet close-up. dorsal (left), ventral (middle), and lateral (right) views F-H: Cryptantha scoparia F: Herbarium specimen G: Fruit, showing smaller calyx H: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. 44 Crossosoma 35(1). Spring-Summer 2009 CTmuricata var. muni cat aj C. muricata var ytjorwsii C. muricata^ &wwnesi ij K?. muricjatal Kar xionc. 1 4 nutlets.per nutlets lance-ovate, coarsely tuberculate no dorsal ridge intermedia C. intermedia Gi m intertneaia 4 nutlets per fruit pJC. muricata D : a ar. denticulata v Figure 22. A-F: Cryptantha muricata. A: C. muricata var. jonesii. Herbarium specimen, showing erect, primary axis bearing numerous lateral inflorescences. B: C. muricata var. denticulata. Herbarium specimen, showing numerous ascending secondary branches and absence of strong primary axis C: C. muricata var. muricata Flower close-up, showing large corolla. D-F: C. muricata var .jonesii D: Corolla, small E: Fruit, with 4 attached nutlets. F: Nutlet close- up, dorsal, ventral, and lateral views. Note ovate shape and dorsal ridge G-l: Cryptantha intermedia var. intermedia. G: Flower with large corolla. H: Fruit, sepals removed, showing four nutlets. I: Nutlet close-up, dorsal, ventral, and lateral views. Note absence of dorsal ridge. Crossosoma 35( 1 ), Spring-Summer 2009 45 49. Cryptantha nmricata (Hook. & Am.) A. Nels. & J.F. Macbr. var. muricala (Figure 22C, p.44) Common name: Showy prickly cryptantha. Distribution in Southern California: Southwestern California region (Transverse and Peninsular Ranges): Imperial. Kern. Los Angeles, Orange. Riverside, San Bernardino. San Diego, Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: This variety is distinguished from the others by the wider (3-8 mm) corolla and grayish nutlets. 50. Cryptantha intermedia (A. Gray) Greene var. intermedia (Figure 22G-I. p.44) Common name: Common/Nievitas cryptantha. Distribution in Southern California: Southwestern California region (South Coast, Channel Islands - San Clemente and Santa Catalina Islands. Transverse and Peninsular Ranges): Imperial. Inyo, Kern. Los Angeles, Orange, Riverside, San Bernardino. San Diego. Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: Corolla relatively wide (3-6 mm), showy: nutlets 4 per fruit, homomorphic, round-margined, lance-ovate to ovate, tuberculate, lacking a dorsal ridge. Note: C. intermedia var. grandiflora [=C. hendersonii (Nelson) Piper], not native to Southern California, has much larger corollas. 51. Cryptantha barbigera (A. Gray) Greene var. fergusoniae J.F. Macbr. (Figure 23A-E. p.46) Common name: Palm Springs cryptantha. Distribution in Southern California: western Colorado Desert, especially Coachella Valley; Imperial, Riverside. San Bernardino San Diego counties. Conservation Status: none. Diagnosis: This variety is largely limited in distribution to the Coachella Valley and periphery. It is distinguished from C. intermedia by the significantly larger (4-6 mm in flower. 5-10 mm in fruit) calyx and from C. barbigera var. barbigera by the larger (4-9 mm wide) corolla. [C. barbigera (A. Gray) Greene in part , sensu W.A. Kelley and Wilken 1993] 52. Cryptantha traskiae I.M. Johnst. (Figure 23F-H, p.46) Common name: Trask's cryptantha. Distribution in Southern California: Southwestern California region (Channel Islands - San Clemente. San Nicolas Islands): Los Angeles, Ventura counties. Conservation Status: CNPS List IB. 2. Diagnosis: Corolla small (1-2 mm wide), nutlets round-margined, lance-ovate to ovate, tuberculate. 1-1. 5(1. 8) mm long, lacking a dorsal ridge. 53. Cryptantha simuians Greene (Figure 24A-C. p.47) Common name: Pine cryptantha. Distribution in Southern California: Southwestern California region (Transverse and Peninsular Ranges); Kern, Los Angeles. Riverside. San Bernardino, San Diego. Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis: Corolla small (1-2 mm wide); nutlets 4 per fruit, round-margined, ovate with dorsal face flattened to low-convex, densely papillate - sparsely tuberculate, ca. 2 mm long, lacking a dorsal ridge. 46 Crossosoma 35(1), Spring-Summer 2009 &fnwbigera var W^Jjglls°niae , C^naiini zeiiaA ,v '^Je^gusomqe\ wa Y+jej^iscmiae Figure 23. A-E: Cryptantha harbigera var. fergusoniae. A: Whole plant from herbarium sheet. B: Fruit C: Close-up of stem (center) and leaf (upper right). D: Flower, showing large corolla. E: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. F-H: C. traskiae. F: Whole plant from herbarium specimen, showing short plant stature G: Inflorescence. H: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. Crossosoma 35( 1 ), Spring-Summer 2009 47 C> simulant C. simulansj [fcfT lokTyij. C. clokeyi C. clokeyi nutlet tubercles translucent iK ; \ «• C. si mu Ians Figure 24. A-C: Cryptantha simulans. A: Whole plants from herbarium specimens. B: Fruit close-up C: Nutlet close-up. dorsal (left), ventral (middle), and lateral (right) views. D-l: Cryptantha clokeyi. D: Whole plant from herbarium sheet E: Close-up of stem (right) and leaf (far left). F: Flower close-up. Note small corolla. G: Fruit H: Nutlet close-up. dorsal (left), ventral (middle), and lateral (right) views. I: Nutlet close-up. showing rounded, translucent tubercles. 54. Cryptantha clokeyi I.M. Johnst. (Figure 24D-1. p.47) Common name: Clokey’s cryptantha. Distribution in Southern California: Mohave Desert; Inyo, Los Angeles, San Bernardino counties. Conservation Status: CNPS List 1B.1. Diagnosis: Corolla small (1-2 mm wide); nutlets 4 per fruit, ca. 3 mm long, round-margined, ovate, densely tuberculate with tubercle tips translucent, lacking a dorsal ridge. 48 Crossosoma 35(1). Spring-Summer 2009 'tlbigenaMawly. ^^a^b igej&var. b. nutlet sculpturing coarsely tuberculate t . to muricate, %' * jtw not papillate \ %* C. bar big era var. barbigera Figure 25. A-C: Cryptantha echinella A: Whole plant from herbarium specimen B: Inflorescence. C: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views D-H: C. barbigera var. barbigera D: Whole plant in field. E: Inflorescence close-up. Note small corollas F: Fruit, showing long calyx G: Nutlet close-up, dorsal (left) and ventral (right) views H: Flower, show ing small corolla Crossosoma 35(1), Spring-Summer 2009 49 55. Cryptantha echinella Greene (Figure 25 A-C, p.48) Common name: Spiny cryptantha. Distribution in Southern California: Tehachapi mountains. Southwestern California region (Transverse Ranges), Mohave Desert; Inyo. Kern, Los Angeles, Mono, San Bernardino. Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Calyx 5-6 mm long in fruit; corolla small (1-1.5 mm wide); nutlets 4 per fruit, ca. 1.5-2 mm long, round- margined, ovate, finely tuberculate and papillate, lacking a dorsal ridge. 56. Cryptantha barbigera (A. Gray) Greene var. barbigera (Figure 25D-H. p.48) Common name: Bearded cryptantha. Distribution in Southern California: Desert province. Tehachapi mountains; Imperial, Inyo, Kern. Los Angeles, Riverside, San Bernardino, San Diego, Santa Barbara. Ventura counties. Conservation Status: none. Diagnosis: Calyx in fruit long (5-10 mm); nutlets 1-4 per fruit, 1.5-2 mm long, round-margined, ovate, densely tuberculate-muricate, lacking a dorsal ridge. This variety is distinguished from C. b. var. fergusoniae by the relatively small corolla (1-2.5 mm wide). [C. barbigera , in part, sensu W.A. Kelley and Wilken 1993] NUTLET COMPARISON WITHIN MAJOR GROUPS For comparison of nutlets within major groups, see Figures 26-30 (at end of article) for images of all Southern California Cryptantha taxa, arranged as follows: Oreocarva Group (Figure 26), Krynitzkia Group, smooth nutlets (Figure 27), Krymitzkia Group, nutlets with ridged or winged margin (Figure 28), Greeneocharis Group (Figure 29A,B), Eremocarya Group (Figure 29C,D), Krynitzkia Group, heteromorphic nutlets with rounded margin (Figure 29 E-H). Krynitzkia Group, rough nutlets with rounded or sharp-angled margin (Figure 30). CONSERVATION STATUS AND ECOLOGY OF SOUTHERN CALIFORNIA CRYPTANTHA Nine species of Southern California Cryptantha are included in the 2009 California Native Plant Society (CNPS) Inventor}7 as sensitive or rare taxa. Of these, two species, Cryptantha clokeyi and C. ganderi, are on CNPS List 1B.1, which signifies that they are “rare, threatened, or endangered in California and elsewhere” and “seriously endangered in California (over 80% of occurrences threatened/high degree and immediacy of threat)/' Cryptantha clokeyi is found in “sandy or gravelly soil” (R.B. Kelley in prep) in the Mohave Desert region, in three counties (Inyo, Los Angeles, and San Bernardino). Only 16 original collections are cited in the Consortium of California Herbaria at the time of this writing. Cryptantha ganderi is found in “stabilized, +- silty, fine sand deposits, creosote scrub association" (R.B. Kelley, in prep.), solely in parts of the Borrego Valley of San Diego County. Only 9 original collections are currently cited in the Consortium of California Herbaria. 50 Crossosoma 35(1). Spring-Summer 2009 Tw o Southern California Cryptantha species are on CNPS List 1B.2. meaning “rare, threatened, or endangered in California and elsewhere” and “fairly endangered in California (20-80% occurrences threatened).” One of these, C. roosiorum , is a distinctive member of Oreocarya. It is known from “rocky, silty soils, dry meadows in open bristlecone pine-limber pine forest, limestone endemic” (R.B. Kelley, in prep.) of only three nearby localities in the Inyo Mountains (Inyo County). Only 5 original collections are cited in the Consortium of California Herbaria. The second CNPS List 1B.2 species is C. traskiae , which is found only on two of the Channel Islands: San Clemente Island (Los Angeles County) and San Nicolas Island (Ventura County), both of the southern group. However. C. traskiae has been collected considerably more, as 60 original collections (almost two-thirds by S. A. Junak) are cited in the Consortium of California Herbaria. The five other Cryptantha species included in the CNPS Inventory - C. costata, C. glomeriflora, C. holoptera, C. scop aria, and C. tumulosa — are on List 4.3. meaning they are relatively rare, of “limited distribution (watch list)” and are “not very endangered in California (<20% of occurrences threatened or no current threats known).” Cryptantha taxa are found in a variety of substrate types and habitats. Most members of the Eremocatya . Greeneocharis , and Krynitzkia groups grow in sandy or gravel soil. Cryptantha flavoculata, C. hoffmannii. C. humilis, C. roosiorum , C. tumulosa. and C. virginensis of the Oreocarya group and C. inaequata of the Krynitzkia group occur occasionally or solely in limestone-based substrates. Interestingly, members of the perennial Oreocarya group tend to occur in high elevation, mountainous regions; the taxa in Southern California are found at elevations ranging from 840-3600 meters (2.700-11.800 feet), w ith a median elevation of 2.300 meters (7,700 feet). In addition, tw o common Cryptantha species - C.flaccida and C. intermedia (no variety indicated) - are cited as “weak” or “indifferent” indicators of ultramafic substrates (Safford et al. 2005). TAXONOMIC PROBLEMS IN CRYPTANTHA Several Cryptantha taxa of Southern California warrant further studies, both quantitative statistical studies of morphology and genetic studies at the population level. Comparative studies of biogeography, ecology, and reproductive biology of the Greeneocharis species pair (C. circumscissa and C. similis ) are needed to better understand how they may be related to one another and how they diverged. Within the Krynitzkia group, the varieties of C. clevelandii need to be properly documented with respect to biogeographic distribution, habitat differences, and variation. Ciyptantha angustifolia is reported to have variants, including forms with homomorphic and winged nutlets, in contrast to the typical heteromorphic. round-margined nutlets, and forms differing in corolla width and vestiture. The complex of varieties in C. muricata are at times difficult to identify and appear to intergrade, particularly in the southern part of the range. Additional research is needed on this group (R. B. Kelley, pers. comm. 2008). What is called C. intermedia var. intermedia shows considerable Crossosoma 35( 1 ), Spring-Summer 2009 51 variation in leaf morphology from coastal to montane habitats. Cryptanthci corollata shows considerable variation. Cryptantha maritima might be worthy of further study with regard to variation in calyx trichome morphology, given that the synonomized C. in. var. pilosa presumably differs in that regard. Both Cryptantha maritima and C. circumscissa would be fruitful to study with respect to potential differences between populations in North versus South America and the assessment of the direction and timing of their dispersal events. In general, it is likely that many collections of Cryptantha across California need careful annotation, using the taxonomy and keys in the upcoming The Jepson Manual (R.A. Kelley, in prep). Just before this article went to press, we received notification of a population of Ciyptantha fendleri (not previously known in California) collected this year in the Little Cowhorn Valley of the Inyo Mountain range, thus technically part of the Southern California region. Cryptantha fendleri. a member of the smooth nutlet Krynitzkia group, will be included in the second edition of The Jepson Manual (R. A. Kelley, in prep) and is currently being considered for CNPS List 2.3 ["rare, threatened, or endangered in California, but more common elsewhere; not very endangered in California (<20% of occurrences threatened or no current threats known)”]. Finally, Southern California Cryptantha that are potentially new to science are currently being evaluated (Jim Andre, personal communication; authors of this paper), which could add even more to the Southern California region's diversity of this fascinating group. ACKNOWLEDGEMENTS We are greatly indebted to the herbaria at COLO, RSA-POM, SD, SDSU, and UC- JEPS for specimen loans, to all the herbaria that are members of the Consortium of California Herbaria (CAS-DS. CDA. CHSC. DAV. HSC, IRVC. OBI. PGM. RSA-POM, SBBG, SD. SDSU, SJSU, UC-JEPS, UCR UCSB. and UCSC, as of this writing) for access to their data, and to UC-JEPS in particular for creating and maintaining the Consortium data interface. We thank Ron B. Kelley for an early version of his descriptions and keys of Ciyptantha . to be incorporated in the second edition of The Jepson Manual. We thank Anna Schochenmaier for photographing several of the high resolution images of nutlets. We thank Gary Monroe for allowing us to use field images of C. cinerea var. abortiva, C. confert {flora, and C. flavoculata. We sincerely thank Andy Sanders and Leroy Gross for reviewing this paper and giving useful suggestions (but of course we take responsibility for any errors). Finally, we thank the editors of Crossosoma for inviting us to publish this article, which is an offshoot of a talk given at the 2008 Southern California Botanists meeting. LITERATURE CITED Abrams, L. 1951. Illustrated flora of the Pacific States. Vol. III. Stanford University Press, Stanford, California. Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society > 141 :399-436. 52 Crossosoma 35(1), Spring-Summer 2009 Buys. M. H. and H. H. Hilger. 2003. Boraginaceae cymes are exclusively scorpioid and not helicoid. Taxon 52:719-724. California Native Plant Society (CNPS). 2009. Inventory of Rare and Endangered Plants (online edition. v7-09b). California Native Plant Society. Sacramento. California. Consortium of California Herbaria. 2009. Data provided by the participants of the Consortium of California Herbaria, http://ucjeps.berkeley.edu/consortium. Don. G. 1836. A General System of Gardening and Botany: Founded upon Miller's Gardener's Dictionary , and Arranged according to the Natural System, volume 4. London, UK. Engler. A., and K. Prantl. 1 897. Die naturlichen Pflanzenfamilien. Engelmann, Leipzig, Germany. Gottschling. M.. H. H. Hilger. and N. Diane. 2001. Secondary structure of the ITS1 transcript and its application in a reconstruction of the phylogeny of the Boraginales. Plant Biology 3:629-636. Greene. E. L. 1887a. Some west American Asperifoliae I. Pittonia 1: 8-23. Greene, E. L. 1887b. Some west American Asperifoliae II. Pittonia 1: 55-60. Greene. E. L. 1887c. Some west American Asperifoliae III. Pittonia 1: 77-120. Hasenstab, K. 2009. Phylogenetic systematics of the genus Cryptantha (Boraginaceae). Master of Science Thesis. San Diego State University, San Diego. California. Heywood. V. H.. R. K. Brummitt, A. Culham. and O. Seberg. 2007. Flowering Plant Families of the World. Royal Botanic Gardens. Kew, Richmond. Surrey. Hickman. J. C. ( ed. ). 1993. The Jepson Manual : Higher Plants of California. University7 of California Press. Berkeley and Los Angeles. Higgins, L. C. 1971. A revision of Cryptantha subgenus Oreocarya. Brigham Young University Science Bulletin Biological Series 8: 1-62. Johnston. I. M. 1925. The North American species of Cryptantha. Dissertation: The Gray Herbarium of Harvard University. Cambridge, Massachusetts. Johnston. 1. M. 1927. A revision of the South American Boraginoideae. Contributions of the Gray Herbarium Harvard University 78:2-1 18. Kelley. R. B. Cryptantha. In The Jepson Manual: Higher Plants of California , second edition, eds. Bruce G. Baldwin, Steve Boyd. D. J. Keil. R. W. Patterson. T. J. Rosatti. and D. H. Wilken. University of California Press, Berkeley. In prep. (Note: title and editors subject to change.) Kelley. W. A., and D. Wilken. 1993. Cryptantha. In The Jepson Manual: Higher Plants of California, ed. J. C. Hickman, 369-378. University of California Press, Berkeley. Mabberley. D. J. 2008. Mabberley's Plant-Book: A Portable Dictionary of the Higher Plants. Their Classification and Uses. 3rd edition. Cambridge University Press. Cambridge. Munz. P. A. and D. D. Keck. 1959. A California Flora. University of California Press, Berkeley Munz. P. A. and D. D. Keck. 1968. Supplement to A Flora of California. University of California Press, Berkeley. Crossosoma 35( I ), Spring-Summer 2009 53 Munz, P. A. 1974. A Flora of Southern California. University of California Press, Berkeley. Payson, E. B. 1927. A monograph of the section Oreocarya of Cryptantha Annals of the Missouri Botanical Garden 14:21 1-358. Reiche, K. 1915. Estudios Criticos de la Flora de Chile. Anales de Universidad de Chile. Santiago, Chile. Rosatti, T. J. 2003. Jepson Online Interchange for California Floristics. Fremontia 31: 23-29. Safford, H. D., J. H. Vires, and S. P. Harrison. 2005. Serpentine Endemism in the California Flora: A Database of Serpentine Affinity. Madrono 52:222-257. Schwarzer, C. 2007. Systematische Untersuchungen an den peruanischen Vertretem der Gattungen Pectocaiya D.C. ex Meisn., Amsinckici Lehm., Plagiobothrys Fisch. & C.A. Mey. und Cryptantha Lehm. ex G. Don (Boraginaceae). Dissertation: Biology Institut fur Biologie Systematische Botanik und Pflanzengeographie, Freie Universitat Berlin, Germany. Steven, P. 2001 onwards. Angiosperm Phytogeny Website. Version 9, June 2008 http: //www. mobot. org/MOBOT/research/APweb Weber, W. A. 1987. Colorado Flora: Western Slope. Colorado Associated University Press. Boulder, Colorado. Wiggins, I. L. 1980. Flora of Baja California. Stanford University Press, Stanford, California. Zuloaga, F., O. Morrone, and M. Belgrano (eds). Catalogo de las Plantas Vasculares del Cono Sur ( Checklist of the Southern Cone), volumes 1-3. Boraginaceae, Cryptantha treatment. In prep. 54 Crossosoma 35(1). Spring-Summer 2009 Figure 26. Oreocarya group nutlets (dorsal view). A: Cryptantha cinerea var. abortiva. B: C. confertiflora C: C. flavoculata. D: C. hoffmannii. E: C. Im mil is. F: C. mi bigen a. G: C. roosiorum. H: C. tumulosa. I: C. virginensis. All images to scale. Cross os oma 35( 1 ), Spring-Summer 2009 55 Figure 27. Krynitzkia group, smooth nutlets (dorsal view). A: Crypt ant ha affinis. B: C. clevelandii var. clevelandii. C: C. clevelandii var. florosa. D: C. flaccida. E: C. gander i F: C. glomeriflora. G: C. gracilis. H: C. leiocarpa. I: C. niaritima. J: C. microstachys. K: C. mohavensis. L: C. ne mad a da M: C. sparsiflora N: C. tor rev ana. O: C. watsonii All images to scale. 56 Crossosoma 35(1). Spring-Summer 2009 Figure 28. Krynitzkia group, nutlets with a ridged or winged margin (dorsal view). A: Cryptantha costata. B: C. holoptera. C: C. inaequata. D: C. oxvgona. E: C. pterocarya xar. pterocarya, winged nutlet (left), unwinged (right). F: C. pterocarya var . purpusii. winged nutlet. G: C. racemosa. H: C. utahensis. All images to scale Crossosoma 35( 1 ), Spring-Summer 2009 57 Figure 29. A-B: Greeneochciris group nutlets (dorsal view). A: Cryptantlia circumscissa. B: C. similis. C-D: Eremocarya group nutlets (dorsal view). C: C. lepida. D: C. micrantha. E-H: Kiynitzkici group, heteromorphie nutlets with a rounded margin (dorsal view, except large nutlet of C. aligns tifolia in lateral view) E: C. angustifolia. F: C. dumetorum. G: C. maritima. H: C. micromeres. All images to scale. 58 Crossosoma 35(1). Spring-Summer 2009 Figure 30. A lynitzkia group, rough nutlets with a rounded or sharp-angled margin (dorsal view ). A: Cryptantha barbigera var. barbigera. B: C. barbigera var . fergusoniae. C: C. clokeyi. D: C. coroll at a. E: C. decipiens. F: C. celt in ell a. G: C. intermedia var. intermedia. H: C. muricata var. denticulata. I: C. mu ri cat a var. jonesii. J: C. muricata var. muricata. K: C. nevadensis var. nevadensis. L: C. nevadensis var. rigida. M: C. recurvata. N: C. scoparia. O: C. simulans. P: C. traskiae All images to scale. Crossosoma 35( 1 ), Spring-Summer 2009 59 Appendix 1. Voucher information for Cryptantha specimens examined and photographed in this study. Cryptantha a/finis (A Gray) Greene: Figure 8A. 27A (SDSU 5410), Cryptantha angustifolia ( Torrey) Greene: Figure 17G.H (SDSU 18676), Figure 171 (SDSU 13030), Figurel7J, 29E (SDSU 5376), Cryptantha barbigera (A Gray) Greene var barbigera Figure 24D,E (SDSU 18686), Figure 24F.H (SDSU 9986), Figure 24G, 30A (SDSU 5378), Cryptantha barbigera (A. Gray) Greene var fergusoneae J.F. Macbr: Figure 25A-C,E, 30B (RSA 712692). Figure 25D (RSA 713328), Cryptantha cinerea (Greene) Cronquist var abortiva (Greene) Cronquist: Figure 5F-H, 26A (RSA 46663), Cryptantha circtimscissa (Hook & Am.) I.M. Johnst. var circuniscissa Figure 3A.B (SDSU 18697), Figure 3C-E, 29A (SD 28881); Cryptantha clevelandii Greene var clevelandii Figure 10F,G (SDSU 14050), Figure 10H. 27B (SDSU 14059), Cryptantha clevelandii Greene var florosa I.M. Johnst : Figure 1 1 A (SDSU 18342). Figure 11B-D, 27C (SDSU 5417); Cryptantha clokeyi I.M Johnst.: Figure 23G-L, 30C (RSA 665270), Cryptantha confertiflora (Greene) Payson: Figure 5B, 26B (RSA 196047), . Cryptantha corollata (I.M. Johnst.) I.M. Johnst Figure 20A.B, 30D (POM 368220), Cryptantha costata Brandegee Figure 16F.H-J (SD 113106). Figure 16G (SDSU 17342), Figure 16K. 28 A (SD 113307); Cryptantha decipiens (M E Jones) A A Heller: Figure 19F,H, 30E (SDSU 18645). Figure 19G(SDSU 16365); Cryptantha dumetorum (A Gray) Greene Figure 18A-D (SDSU 18694). Figure 18E-G, 29F (RSA 553891 ): Cryptantha echinella Greene Figure 24A-C. 30F (RSA 682890), Cryptantha flaccida (Lehmann) Greene: Figure 8D(SD 120625), Figure 8E, 27D(SDSU 17311), Cryptantha flavoculata (A. Nels.) Payson. Figure 5D, 26C (SD 21927) ; Cryptantha ganderi I.M. Johnst : Figure 9C.D, 27E (SD 115050); Cryptantha glomeriflor a Greene: Figure 8B, 27F (SD 87724). Figure 8C (RSA 625075); Cryptantha gracilis Osterh Figure 10A (RSA 709650), Figure 10B. 27G (SD 4205); Cryptantha hoffmannii I M. Johnst : Figure 7A,C (SD 81222), Figure 7B (SDSU 5495). Figure 7D, 26D (RSA 79771), Cryptantha holoptera (A Gray) J.F Macbr Figure 13A.B. 28B (SD 120959), Figure 13C (SDSU 13036); Cryptantha humilis(A Gray) Payson: Figure 6E (SD 87618), Figure 6F, 26E (RSA 516990); Cryptantha inaequata I.M. Johnst Figure 17A-F, 28C (RSA 173141); Cryptantha intermedia (A Gray) Greene var intermedia Figure 22G (SDSU 16074), Figure 22H.I, 30G (SDSU 14564); Cryptantha leiocarpa (Fisher & C. A. Meyer) Greene: Figure 9E,G, 27H (SDSU 19212); Cryptantha lepida (A. Gray) I.M Johnst : Figure 4A-C (SDSU 17572). Figure 4D-E. 29C (SDSU 17281 ), Cryptantha nuiritima (Greene) Greene var maritima Figure 9F.H(SDSU; inset SD 113111). Figure 19A-C(left), 271, 29G (right)(SD 113111), Figure 19C(right). 29G (left) (SDSU 5460), Cryptantha micrantha (Torrey) I.M. Johnst : Figure 4F (SDSU 5388), Figure 4G, 29D (SDSU 5421), Figure 4H,I (SDSU 5431 ). Figure 4J (SDSU 5419); Cryptantha micromeres (A. Gray) Greene: Figure 18H,J (SDSU 16695), Figure 18K (SDSU 15465), Figure 18L-M. 29H (SDSU 12173); Cryptantha microstachys (A. Gray) Greene: Figure 10CJD (SD 58176), Figure 10E, 27J (SD 44966); Cryptantha mohavensis (Greene) Greene: Figure 12A. 27K (SD 129726). Figure 12B (RSA 712528); Cryptantha muricata (Hook & Am.) A Nels & J.F. Macbr var denticulata (Greene) I.M. Johnst Figure 22B. 30H (SD 161808). , Cryptantha muricata (Hook. & Am.) A. Nels & J.F Macbr var. jonesii (A Gray) I.M Johnst.: Figure 22A,E (SDSU 16714), Figure 22D (17512). Figure 22F, 301 (SDSU 5413), Cryptantha muricata (Hook & Am.) A. Nels. & J F Macbr var muricata Figure 22C. 30J (SDSU 19186); Cryptantha nemaclada Greene: Figure 9A,B. 27L (RSA 195150); Cryptantha nevadensis A Nels & PB Kennedy var nevadensis Figure 20C-H, 30K (SDSU 13032); Cryptantha nevadensis A Nels. & PB Kennedy var rigida I.M Johnst Figure 21A-E, 30L (RSA 580318), Figure 21B (POM 287290); Cryptantha nubigena (Greene) Payson: Figure 6C (JEPS 90327), Figure 6D, 26F (SD 87723); Cryptantha oxygona (A. Gray) Greene: Figure 16A-D (RSA 717219), Figure 16E, 28D (RSA 709140); Cryptantha pterocarya (Torrey) Greene var cycloptera (Greene) J.F Macbr Figure 15C (SDSU 17298); Cryptantha pterocarya (Torrey) Greene var pterocarya Figure 14E (SDSU 18692), Figure 14F.G (SDSU 12435), Figure 15A, 28C (SDSU 5500), Figure 15B (SDSU 17304); Cryptantha pterocarya (Torrey) Greene var purpusii Jeps.: Figure 15 D,E, 28F (SD 91861); Cryptantha racemosa (S. Watson) Greene: Figure 13D (SDSU 5494). Figure 13E.G, 28G (SD 42216), Figure 13F (SDSU 18710); Cryptantha recurvata Coville: Figure 19D (RSA 370268). Figure 19E, 30M (RSA 250147), Cryptantha roosiorum Munz Figure 6A.B, 26G (RSA 346621 ). Cryptantha scoparia Nelson: Figure 21F.G (RSA 641530), Figure 21H. 30N (POM 170650), Cryptantha similis K Mathew & PH. Raven Figure 3F-I (RSA 272392), Figure 3J. 29B (RSA 660941 ), Cryptantha simulans Greene: Figure 23D,E (SD 70318). Figure 23F. 300 (SDSU 5488), Cryptantha sparsiflora (Greene) Greene Figure 8F (SD 40827), Figure 8G. 27M (RSA 191197); Cryptantha torreyana ( A. Gray) Greene: Figure 1 1E-I (JEPS 102382), Figure 11 J, 27N (SD 129490); Cryptantha traskiae I.M Johnst : Figure 23A.B (JEPS 93692), Figure 23C, 30P (SD 90374); Cryptantha tumulosa (Payson) Payson: Figure 7H (SD 107276). Figure 71, 26H (SD 58351), Cryptantha utahensis (A Gray) Greene: Figure 14A (SDSU 18688). Figure 14B-D, 28H (SD 100153); Cryptantha virginensis (ME Jones) Payson: Figure 7E (SD 107269), Figure 7F (SDSU 5501), Figure 7G. 26F (RSA 340713); Cryptantha watsonii (A. Gray) Greene: Figure 12C (COLO 503943). Figure 12D, 270 (RSA 51987) Become a Member of Southern California Botanists ] Individual (family) $15.00 | | New Member ] Organization $25.00 | | Renewal Name Address 1 Address 2 City State Zip Code Phone (optional) Email address In addition, I would like to give $ to help support SCB. Make Checks payable to Southern California Botanists and mail to: Treasurer, Southern California Botanists Rancho Santa Ana Botanic Garden 1500 North College Avenue Claremont, C A. 91711 Southern California Botau. - Founded 1927 - New York Botanical Garden Libra 5185 00268 28 http://www.soca/bot. org Membership, Subscriptions, and Back Issues Individual and Family Memberships in SCB are $15 per calendar year domes- tic, 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 foriegn insti- tutions. Back issues (Volume 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. Available SCB Special Publications No. 1 A Flora of the Santa Rosa Plateau , by Earl W. Lathrop and Robert F. Thome, 30 pp $7.00 No. 3 Endangered Plant Communities of Southern California , Proceedings of the 15th Annual SCB Symposium, edited by Allan A. 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Index to Cryptantha of Southern California TEXT IMAGES KEY to Cryptantha Groups p. 10 C. affinis p.21 22 C. angustifolia p. 35.. 37 C. barbigera var. barbigera p. 49 48 C. barbigera var. fergusoneae p. 45 .. 46 C. cinerea var. abortiva p. 15 16 C. circumscissa var. circumscissa P 11- 12 C. clevelandii var. clevelandii p. 25 26 C. clevelandii var. fiorosa p. 25 .. 27 C. clokeyi p. 47 47 C. confertiflora P 15 - 16 C. corollata p. 38 41 C. costata P- 35 .. 36 C. decipiens p. 38 40 C. dumetorum p. 38 .. 39 C. echinella p. 49 48 C.flaccida P-21.. 22 C. flavoculata p. 15 16 C. ganderi p. 23 .. 24 C. glomeriflora P-21 22 C. gracilis p. 23 .. 26 C. hoffmannii p. 17 19 C. holoptera P-31.. 32 C. humilis p. 17 18 C. inaequata p. 35 .. 37 C. intermedia var. intermedia p. 45 44 C. leiocarpa p. 23 .. 24 C. lepida p. 13 14 C. maritima var. maritima p. 23 .. 24, 40 P. micrantha p. 13 14 C. micromeres p. 38.. 39 C. microstachys p. 25 26 C. mohavensis p. 25 .. 28 C. muricata var. denticulata p. 42 44 C. muricata var. j ones ii p. 42 .. 44 C. muricata var. muricata p. 45 44 C. nemaclada p. 23 .. 24 C. nevadensis var. nevadensis p. 38 41 C. nevadensis var. rigida p. 42 .. 43 C. nubigena P- 17 18 C. oxygona p. 35 .. 36 C. pterocarya var. cycloptera p. 35 34 C. pterocarya var. pterocarya p. 31 .. 33, 34 C. pterocarya var. purpusii P-31 34 C. racemosa P-31 .. 32 C. recurvata p. 38 40 C. roosiorum P- 17- 18 C. scoparia p. 42 43 C. similis P- 11 - 12 C. simulans p. 45 47 C. sparsiflora p. 23 .. 22 C. torreyana p. 25 27 C. traskiae p. 45 .. 46 C. tumulosa P- 17 19 C. utahensis P- 31 .. 33 C. virginensis p. 17 19 C. watsonii P- 29 .. 28 Front Cover: Nutlet dorsal images of all 49 species and 56 taxa (including varieties) of Cryptantha in Southern California. All images to scale. pep CROSSOSOMA Journal of the Southern California Botanists , Inc . Volume 35, Number 2 Fall-Winter 2009 Southern California Botanists, Inc. -Founded 1927 - http.V/www.socalbot. org CROSSOSOMA (ISSN 0891-9100) is published twice a year by Southern Cali- fornia Botanists, Inc., a California nonprofit organization of individuals devoted to the study, conservation, and preservation of the native plants and plant com- munities of southern California. SCB Board of Directors for 2009 President Gary Wallace Vice President Naomi Fraga Secretary. Carrie Kiel Treasurer. Alan P. Romspert Webmaster Naomi Fraga Editors of Crossosoma Scott D. White and Michael Honer Editor of Leaflets Kerry Myers Directors-at-large David Bramlet Gilberto Ocampo Sara Baguskus Linda Prince Elizabeth Delk Gina Richmond Erica Gardner Fred Roberts Charlie Hohn Darren Sandquist Diane Menuz Allan Schoenherr Orlando Mistretta Jonathan Snapp-Cook Bart O’Brien Sula Vanderplank and Katie Vinzant Ex officio Board Member Sula Vanderplank (Past President) Articles, book reviews, or other items for submission to CROSSOSOMA can be sent to the editor Scott White (scottbioservices@verizon.net) or 201 N. First Ave., #102, Upland, CA., USA, 91786. Electronic submission is preferred. Please see our website, www. socalbot.org, for format guidelines. Notices of a time-dated nature (field trips, work- shops, symposia, etc.) to be included in the newsletter Leaflets should be submitted to Kerry Myers, Editor of Leaflets , kerrymyers@fs.fed.us, or mail to: Kerry Myers, Botanist, SBNF Mountaintop Ranger Dist., 42300 North Shore Dr., Fawnskin, CA 92333-04004. 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 specifically stated. Copyright © 2009 by Southern California Botanists, Inc. All rights reserved. Permission to reproduce items in CROSSOSOMA, in whole or part, should be requested from the Editor. Crossosoma Volume 35, Number 2 Fall- Winter 2009 Published February 2010 CONTENTS DEDICATION: Alan Romspert, 1945-2009 Allan Schoenherr 61 Lichens and Lichenicolous Fungi of the Northwestern Santa Ana Mountains Kerry Knuds en and Jana Kocourkova 66 New Records of Lichens and Lichenicolous Fungi for California I. Jana Kocourkova, Kerry Knuds en and Othmar Breuss 82 Cost and Effectiveness of Small-Scale Foeniculum vulgare Control Methods Abigail Gw inn Noteworthy Collection .87 96 Cover: A saxicolous lichen community on sandstone in Weir Canyon. In center of pic- ture, Candelariella rosulans (yellow) growing with Caloplaca squamulosa (orange) with Lecanora mural is (light gray) directly to left and right. Photograph by Rolf Muqrtter. 6ui Ftb i b /if m jyi?^rypfpu r yr\m. A1 Romspert in Madagascar, June 2006. Photo: Allan Schoenherr Crossosonui 35(2), Fall-Winter 2009 61 DEDICATION: Alan Romspert (Roms), 1945-2009 One of California's premier desert botanists, Alan Romspert, passed away on Wednesday, August 19, 2009. I will miss him dearly. Roms and 1 traveled and camped together for years. We collected plants, backpacked in the mountains, fished, and explored the southwest deserts. When 1 was still teaching at Fullerton College, I would stay at his house about once a week. We would sit on his front porch, drink whiskey, and solve the world's problems. He would smoke a cigar. It was kind of like Alan Shore (James Spader) and Denny Crane (William Shatner) of the television show ‘‘Boston Legal.” We, of course, earned this ritual to campfires on our many camping trips. I first met Roms in the early 1970s when he was freshly back from Viet Nam. We both took a class in Desert Biology at California State University, Fullerton. He went on to get his Bachelor’s and Master's degrees there. Even though we most often thought of Romspert as a Botanist, he did his early research on the physiology of Amphibians, and he published two scientific papers on water relations in Amphibians. He also coauthored a paper with Jack Burk on the plants of the Algodones Dunes in Imperial County. Alan was a member of Southern California Botanists since the early 1970s. Over the years, he held every office except secretary. He served as Treasurer for the last 30 years, during which time Board members will agree (or perhaps admit) that he was the glue the held the organization together. Alan kept the records, he balanced the books, wrote checks, reported our financial dealings to the IRS and kept up our non-profit status. He printed the mailing labels for Leaflets and Crossosoma , dutifully highlighting the expiration date of errant members. To save a bit of postage, he hand-delivered copies to members he knew. He sorted the mail by zip code and delivered it to the Post Office. He kept the membership records and, like a badger, he pursued members who hadn’t paid their dues on time. Most of Southern California Botanists’ records, materials and archives were kept in boxes at Alan’s house. When it came time to distribute research grants, Alan headed the committee that read the proposals. Roms took care of records and finances his own way. Nobody else understood it, but it always got done. He was a bit set in his ways and rarely saw reason to change them (though he did finally upgrade his ancient membership data base to a system compatible with 21st century computers). He could be a curmudgeon at board meetings, sometimes frustrating the other Directors, but entertaining to at least some (the Directors stifling their laughter or hiding it behind their paperwork). 62 Crossosoma 35(2), Fall -Winter 2009 The Southern California Botanists symposium is our single, important social, scientific, and fund-raising event each year. Most of our members look forward to the long day of botany, and chance to meet new and old friends and colleagues, and maybe meet afterwards for beer. The Symposium is now in its 34th year. Without Romspert’s efforts, the Symposium would never have lasted. Each year, Roms reserved the Ruby Gerontology Center and made sure that campus security opened the doors at 7:00 am so we could start setting up. He kept track of early registrants. He would borrow a pickup truck, and sometimes single-handedly transport large folding tables from storage to the symposium site. He made the coffee, heated water, set up cold drinks, pulled tea bags, sugar, cream, napkins, hot cups, cold cups from storage, and bought doughnuts on the day of the symposium. We then set up tables and displays featuring SCB field trips, publications, and T-shirts. During the speakers’ presentations he often remained outside to continue sales and welcome late-comers. He wrote receipts and tallied up the sales. He also organized and mediated the silent auction where funds for the Susan Hobbs grant were gathered. Romspert’s annual symposium punch is legendaiy. It consisted of the sodas, bottled drinks, and powdered drinks left over from his year’s camping trips, supplemented with ice and a few jugs of fruit punch or fizzy water on clearance at the market. When it was all over, Roms reported to the Board on attendance, sales, and expenses, detailing every line item to the exact penny. In 1976, when the program started, Roms began a 23 year stint with the Desert Studies Center at Zzyzx in the Mojave Desert near Baker. His contributions to the Center are immeasurable. He was instrumental in all aspects of building or restoring the physical facilities, which were in a state of disrepair (they were originally obtained from an evangelist who ran a rehabilitation facility there, on public land, but without the permission of the BLM). As an expert on the desert flora, his knowledge was unrivaled. It was joked that he personally knew all the plants, calling them by name, “Joe,” “Sally,” etc. Through his botanical collecting, specimen preparation, and curation work, he produced for the Center the best herbarium in California's Mojave Desert. He collected extensively in the Panamint Mountains with the idea in mind that ultimately he would publish a flora of that mountain range. The Rancho Santa Ana Botanic Garden Herbarium now houses nearly 600 of his specimens collected in the Panamints. At the Desert Studies Center Roms taught numerous courses sponsored through Cal State San Bernardino and other universities. I shared teaching duties with him numerous times, including recent years when he and I co-taught the “Flora of Joshua Tree National Park,” offered as an extension course through the University of California, Riverside. Crossosoma 35(2), Fall -Win ter 2009 63 Among the groups to which he belonged he always was a leader. He was on the board of directors of the Desert Explorers, where his nickname was “Flower Child." Desert Explorers sponsors 4-wheel drive excursions to desert locations. Roms and I lead together many of their field trips, always incorporating natural history into them. He also organized the highly successful Silent Auction at the Desert Explorer Rendezvous. And he was active in The Ancient and Honorable Order of E Clampus Vitus (the Clampers), a service group that establishes and builds historical monuments throughout the state. Alan's love for the desert, bad roads, remote camps, and quirky sites and celebrations, may have made him an archetypical Clamper. Alan Romspert loading his plant press, Mojave Road, 1992. Photo: Sherry Schmidt We shall be telling Romspert stories for years. He was a skinflint and a collector. Besides plants, he collected bottles of hot sauce, and he had a huge stamp collection. I don’t think he threw anything away. He saved everything including bottle caps. He clipped coupons from the “Penny Saver,” and dutifully submitted receipts and rebate coupons. On desert trips he picked up discarded bottles and aluminum cans which he turned in for refunds. 64 Crossosoma 35(2), Fall -Winter 2009 Many of us remember his “thrifty” ways. He would drive out of his way to save a few pennies on gasoline. One of the great stories goes back a number of years, when Roms was part of a group that was camping in the Avawatz Mountains at Sheep Creek Springs. He had left a pair of his shoes at a microwave tower on the south road to Death Valley. He talked the group into piling into a single vehicle and driving back to retrieve his shoes. It was farther than anyone expected, but he kept reassuring them it was just a little bit farther. As it turns out it was many miles away and took about a half day of driving to retrieve his pair of nearly worn out “flip-flops.” That experience, among his friends, led to a new unit of measurement known as a “Roms,” an unknown distance that is probably farther than you think. “How far is it to Grinderswitch Spring?” “I’m not sure, I’d say it’s about three Roms!!” In spite of his apparent thriftiness and gruff exterior Roms was a caring and generous person. He would loan money to a hapless friend, never expecting to be repaid. He would buy something, usually at a swap meet, and then give it to a friend he thought could use it. Once, knowing I was building a new deck, he bought for me 15 gallons of the old, oil-based formula Thompson’s Water Seal, and never would tell me how much it cost. His friends will remember his famous salsa. About once a year he would buy flats of tomatoes, onions, and chiles and spend two days cooking and canning. While Dave McClanahan most often helped with the process, I will never forget the tear-streaked hours, crying, while 1 chopped onions. Traditionally, at Christmas, he distributed jars of salsa to anyone who would take them. Nearly everyone did. In his honor, I may never open my last jar of salsa. It was in May of 2007 when the specter of cancer reared its ugly head. Roms and I were sitting around a campfire in Catavina. We were on our way back from Cabo San Lucas, traveling in his black Nissan. Roms asked me about a lump under his left ear, covered by his beard. It turned out to be non-Hodgkins lymphoma. He was treated with chemotherapy and radiation and subsequently was cleared of any cancer, but apparently the chemotherapy had taken a toll on his heart. He began to experience shortness of breath and congestive heart failure was diagnosed. He had been receiving treatment for that problem, but the shortness of breath continued and in recent months he lacked energy. It didn’t stop him. He continued his activities including his now famous roll-over on the road to Catavina in 2008, and in his “newest” Nissan he accompanied the Desert Explorers on the Neal Johns trip to Baja in May 2009. In fact, Roms went fishing on Friday, August 14, two days before his long-time girl friend, Linda Harris, took him to the hospital for the last time. Once in the hospital he deteriorated rapidly. Associated with his weak Crossosonui 35(2), Fa] I -Win ter 2009 65 heart and low blood pressure, blood supply to vital organs became diminished, ultimately leading to kidney failure and reduced liver function. Life support was removed about noon on Wednesday and he was gone by 12:15 PM. Alan Romspert was my friend. We traveled, botanized, camped, and hiked together. I shall never forget him, one of the best friends I or anyone could ever have. by Allan Schoenherr Roms at Catavina, Baja California, May 2007. Photo: Allan Schoenherr 66 Crossosoma 35(2), Fall -Winter 2009 LICHENS AND LICHENICOLOUS FUNGI OF THE NORTHWESTERN SANTA ANA MOUNTAINS Kerry Knudsen The Herbarium, Dept, of Botany & Plant Sciences, University of California, Riverside, California 92521 kk999@msn.com Jana Kocourkova Faculty of Environmental Sciences, Czech University of Life Sciences Prague, Kamycka 129, CZ-165 21 Praha 6 - Suchdol, Czech Republic kocourkovaj@fzp.czu.cz ABSTRACT: 169 taxa are reported for the northwestern Santa Ana Mountains, comprising 1 56 lichens and 1 3 lichenicolous fungi. Sarcogyne plicata H. Magn. is removed from synonymy with Sarcogyne privigna (Ach.) A. Massal. KEYWORDS: Biodiversity, California, Fremont Canyon, Floristics, Orange County, Weir Canyon. Fig. 1. Fremont Canyon. Photo: Janet Good. Crossosoma 35(2), Fall-Winter 2009 67 Fig. 2. Weir Canyon sandstone slabs. Photo: Rolf Muertter woodlands, grassland, chaparral, coastal sage scrub and riparian woodlands cover the hills and valleys. Two major canyons, Fremont Canyon (Fig. 1) and Weir Canyon on the Irvine Ranch (Fig. 2), were surveyed in 2006-2008. The elevational range of the study area was approximately 252 meters (827 feet) to 600 meters (1968 feet). INTRODUCTION The Santa Ana Mountains are among the Peninsular Ranges in southern California extending over 64 km (40 miles), roughly from the Santa Margarita River in San Diego County to Santa Ana Canyon in Orange and Riverside counties. The highest peak in the range is Santiago at 1734 m (5,689 ft) (Lathrop & Thorne 1978; Boyd et al. 1995). The known vascular flora of the range consisted of 1044 known taxa in 1995 (Boyd and Roberts 1995). The northwestern end of the Santa Ana Mountains is in Orange County along the Santa Ana River west of Sierra Peak and in the foothills and valleys bordering Santiago Creek on the edge of the coastal plain. Sandstones predominate. Oak 68 Crossosoma 35(2), Fall -Winter 2009 This was the first professional lichen inventory in Orange County. We collected 597 specimens of lichens and lichenicolous fungi, which were curated and databased at the Herbarium of the University of California at Riverside (UCR). Weir Canyon was surveyed in 2006. Fremont Canyon was surveyed in 2007 and 2008 after fire devastated the canyon in early 2006, and this part of the study was twice interrupted by the Weir Canyon fire in February 2007 and the Santiago fire in October 2007. Over 60 days were spent in the field. Collecting was intuitive and subjective. Some days no collections were made due to the devastation caused by the Fremont Canyon fire, which incinerated many acres of all lichens (Fig. 3). Thin-layer chromatography (TLC) was performed on selected specimens by J. C. Lendemer (NY) and J. A. Elix (CANB). Fig. 3. Upper Fremont Canyon. Fire burned lichens off many boulders. Photo: Janet Good. Corticolous or lignicolous lichens occur on bark or wood, even fence posts, and can be specific to certain vascular genera. Saxicolous lichens occur on various rock substrates and some species specifically occur on calcareous or acidic substrates (Fig 4). Terricolous lichens occur on soil. Some lichens are parasitic on other lichens, usually as juveniles, but become eventually independent and are termed lichenicolous lichens. Lichenicolous fungi are non-lichenized microfungi that are symbiotic with lichens and have co-evolved with them and are often host specific Crossosoma 35(2), Fall-Winter 2009 69 Fig. 4. Lecanora muralis , a common saxicolous lichen on sandstone. Photo: Rolf Muertter. on the species or genus level. More information can be found on most species in the three volumes of the Lichen Flora of the Greater Sonoran Desert Region which has excellent but incomplete coverage of Southern California (Nash et al. 2002. 2004, 2007) or in the literature cited in this paper. More detailed information on individual collections is available online at the UCR Herbarium website (http:// sanders5.ucr.edu/lichensflat_index.php). Abbreviations are used for Fremont Canyon (F) and Weir Canyon (W). Frequency is based on subjective observations. 70 Crossosoma 35(2), Fall -Winter 2009 CHECKLIST OF THE LICHENS AND LICHENICOLOUS FUNGI OF THE NORTHWESTERN SANTA ANA MOUNTAINS Acarospora badiofusca (Nyl.) Th. Fr. Saxicolous. Common (F & W). Acarospora fuscata (Schrad.) Arnold - Saxicolous. The authority for this species needs investigation. Rare (W). Acarospora hassei Herre - Saxicolous. Infrequent on small granite pebbles and on sandstone outcrops (W). Acarospora obnubila H. Magn. - Saxicolous. Frequent. (F). Acarospora obpallens (Nyl. ex Hasse) Zahlbr. - Saxicolous, Terricolous. Often densely pruinose. Common (F&W). Acarospora robiniae K. Knudsen- Saxicolous. A coastal species. One population was found on top of a sandstone outcrop (F). Acarospora rosulata H. Magn. - Saxicolous. Treated as A. bullata Anzi, which we no longer recognize as occurring in North America. (Knudsen 2007; Knudsen et al. 2010). Rare (F). Acarospora schleiclteri (Ach.) A. Massal. - Terricolous. Once common in southern California (Hasse 1913). Rare on alluvium on sandstone outcrop on north ridge of Fremont Canyon (F). Acarospora socialis H. Magn. - Saxicolous. Common (F & W). Acarospora terricola H. Magn - Terricolous. Infrequent (F & W). Acarospora the/ococcoides (Nyl.) Zahlbr. — Terricolous. Endemic to California and Baja. Type collected by Orcutt in San Diego. Rare on alluvium on sandstone slabs. (F). Acarospora veronensis A. Massal. - Saxicolous. Infrequent on small stones (W). Aspicilia confusa Owe-Larss. & A. Nordin - Saxicolous. Common gray species, described from Tenaja Canyon in Santa Ana Mountains (F & W). Aspicilia glaucopsina (Nyl. ex Hasse) Hue - Terricolous, Saxicolous. Persisting on sandstone outcrops. Infrequent (F & W). Aspicilia pacifica Owe-Larss. & A. Nordin - Saxicolous. Maritime species. Infrequent (F & W). Aspicilia p/taea Owe-Larss. & A. Nordin - Saxicolous. Only found on small stones scattered in chaparral on ridge of Fremont Canyon (F). Buellia badia (Fr.) A. Massal. - Saxicolous, lichen icolous. Common (F & W). Buellia punctata (Hoffrn.) A. Massal. - Corticolous. Common especially on old or dead branches of Adenostoma fas cicul alum, Salvia mellifera and Sambucus mexicana (F & W). Buellia ryanii Bungartz - Saxicolous. Only found on small stones of granite or rhyolite washed out of the sandstone. Infrequent (F & W). Buellia sequax (Nyl.) Zahlbr. - Saxicolous. Common (F & W). Buellia tesserata Korb. - Saxicolous. Maritime species. Rare (F). Crossosoma 35(2), Fall -Winter 2009 71 Caloplaca a yen arid (Pers.) Mull. Arg. - Saxicolous. Common (F & W). Caloplaca atroflava (Turn.) Mong. - Saxicolous. On concrete. Rare (W). Caloplaca bolacina (Tuck.) Herre - Saxicolous. Maritime species. Frequent (W). Caloplaca citrina (Hoffm.) Th. Fr. - Saxicolous. Wetmore's concept of this species is heterogeneous and includes a common saxicolous leprose taxon found in Riverside County (Wetmore 2007). It is unclear whether this taxon is C. citrina s. str. Caloplaca taxonomists in Europe are skeptical that C. citrina even occurs in North America (Vondrak , pers. comm.) Rare (W). Caloplaca cremtlalella (Nyl.) Oliv. - Saxicolous. Common (F&W). Caloplaca decipiens (Arnold) Blomb. & Forss. - Saxicolous. Frequent (F). Caloplaca epithallina Lynge - Lichenicolous fungus common on saxicolous crustose lichens (F & W). Caloplaca microphyllina (Tuck.) Hasse - Corticolous. Abundant on one Quercus agrifolia (W). Caloplaca /ws/i/i Nav.-Ros., Gaya & Hladun - Saxicolous. Common (F & W). Caloplaca pyracea (Ach.) Th. Fr. - Corticolous. This name is tentatively used in California for those species treated by Wetmore on bark as Caloplaca holocarpa (Wetmore 2007; Arup 2009). Common on Quercus agrifolia , chaparral and old coastal sage shrubs (F&W). Caloplaca saxicola (Hoffm.) Nordin - Saxicolous. Infrequent (F). Caloplaca squamosa (B. de Lesd.) Zahlbr. - Saxicolous. Frequent (F). Caloplaca subsoluta (Nyl.) Zahlbr. - Saxicolous. Rare (W). Candelaria pacifica Westberg - Corticolous, occasionally saxicolous. Common on Quercus agrifolia , chaparral and coastal sage shrubs (F & W). Candelariella aure/la (Hoffm.) Zahlbr. - Saxicolous. Common (F & W). Candelariella rosulans (Miill. Arg.) Zahlbr. - Saxicolous. Infrequent (F). Candelariella vitellina (Hoffm.) Miill. Arg. - Saxicolous. Frequent (F & W). Carhonea latypizodes (Nyl.) Knoph & Rambold- Saxicolous. Common (F & W). Cercidospora caudata Kernst. - Lichenicolous fungus infrequent on apothecia of Caloplaca squamosa (F). Chyrsotrix candelaris (L.) J.R. Laundon - Corticolous. Probably heterogeneous in California and needs to be revised. Abundant on occasional Quercus agrifolia (W). Cladonia acuminata (Ach.) Norrlin - Terricolous. This specimen contained norstictic acid. The specimen was poor and could not be identified positively by morphology and our identification awaits verification by further collections. If positively identified the species would be new state record. (Tucker & Ryan 2006). Rare (W). Cladonia chlorophaea (Florke ex Sommerf.) Spreng. - Terricolous. Infrequent (F & W). 72 Crossosorna 35(2), Fall-Winter 2009 Cladonia fimbriata (L.) Fr. -Terricolous and sometimes lignicolous. Infrequent (W). Cladonia hammeri Ahti - Terricolous. Endemic to southern California and Baja. Infrequent (F & W). Cladonia nashtii Ahti - Terricolous. Common (F & W). Cladonia pyxidata (L.) Hoffm. - Terricolous. Infrequent (F). Cladonia scabriuscula (Delise) Nyl. - Terricolous. Commonly found on detritus beneath chaparral (F & W). Cladonia subfimbriata Ahti - Terricolous. Infrequent. (W). Collema tenax (Sw.) Ach. - Terricolous, sometimes saxicolous on sandstone. Frequent ( W). Dermatocarpon americanum Vain. - Saxicolous. Rare, though often common in southern California (F). Dintelaena radiata (Tuck.) Mull. Arg. - Saxicolous. A dominant maritime species which occurs inland at scattered locations to Bautista Canyon at base of San Jacinto Mountains. Infrequent (F & W). Diploicia canescens (Dicks.) A. Massal. - Corticolous. A maritime species. This is the farthest from coast it has been collected in southern California. Rare on Malacothamnus fasciculatus and Quercus agrifolia. (W). Dip/osch isles actinostomus (Ach.) Zahlbr. — Saxicolous. Common on sandstone on upper ridges of Fremont Canyon (F). Dip/osc/iistes niuscorum (Scop.) R. Sant. - Terricolous, lichenicolous. Common lichen parasitic on Cladonia species (F & W). Diploschistes scruposus (Schreb.) Norman - Saxicolous. Common (F & W). Diplotonnna a/boatruni (Hoffm.) Flot. - Saxicolous. Syn. Buell ia alboatra. The genus placement is still open to question and the species is in need of revision. One locally abundant population on sandstone (F). Endocarpon loscosii Mull. Arg. - Terricolous. Frequent but often sterile, spreading by anastomosing rhizohyphae (F & W). Endocarpon pa/liduluni (Nyl.) Nyl. - Saxicolous. Infrequent (F & W). Endocarpon pusillum Hed. - Terricolous. Common (F & W). Flavoparmelia baltimorensis (Gyel. & Foriss) Hale - Saxicolous. Locally abundant on shaded sandstone slab on a north-facing slope of Weir Canyon (W). Flavoparmelia caperata (L.) Hale - Corticolous. Frequent on Quercus agrifolia (W). F/avopunctelia flaventior (Stirt.) Hale - Corticolous. The most common macrolichen on Quercus agrifolia (F & W). Flavopunctelia soredica (Nyl.) Hale - Saxicolous, corticolous. Rare (W). Fuscopannaria coralloidea F. M. Jorg. - Terricolous. Rare (W). Crossosonui 35(2), Fall -Win ter 2009 73 Gelatinopsis acarosporicola Kocourk. & K. Knudsen -Lichenicolous fungus on Acarosporci socialis described from and currently only known from Fremont Canyon (Kocourkova & Knudsen 2009). Rare (F). Hyperphyscia adglutinata (Florke) H. Mayrh. & Poelt- Corticolous. Abundant on scattered Ouercus agrifolia (F & W). Intralichen baccisporus D. Hawksw. & M.S. Cole - Lichenicolous fungus infrequent on apothecia of Cciloplaca species (F). Lecania brunonis (Tuck.) Herre - Saxicolous. Common (F & W). Lecania cyrtella (Ach.) Th. Fr. - Corticolous. Rare on dead branches of Salvia me 1 1 if era (W). Lecania hassei (Zahlbr.) W. Noble - Saxicolous. Syn. Lecania brattiae. Common (F & W). Lecania naegelii (Hepp.) Diederich & v.d. Boom. - Corticolous. Rare on Malacothamnus fascicul at us ( W) . Lecania toninioides Zahlbr. - Terricolous, rarely saxicolous. Rare on sandstone (F). Lecanora caesiorubella Ach. - Corticolous. Rare on Qurcus agrifolia (W). Lecanora gangaleoides Nyl. - Saxicolous. Frequent (F & W). Lecanora hagenii (Ach.) Ach. - Corticolous, saxicolous. Frequent on sandstone and concrete (F & W). Lecanora munzii K. Knudsen & Lendetner - Corticolous. On dead weathered chaparral wood on ground. Recently described from Claremont (Knudsen & Lendemer 2009). Rare (W). Lecanora muralis (Schreb.) Rabenh. - Saxicolous. Common (F & W). Lecanora pseudistera Nyl. - Saxicolous. Common (F & W). Lecanora simeonensis K. Knudsen & Lendemer - Lignicolous. On wood of Adenostoma fasciculatum. We recently described this species from San Simeon (Lendermer & Knudsen 2009). The thallus was leprose and no fertile specimens were seen. This report extends the range south from Montana de Oro in San Luis Obispo County. Infrequent on chaparral (W). Lecanora subimmergens Vain. - Saxicolous. Common (F & W). Lecidea cinerata Zahlbr. - Saxicolous. Rare California endemic originally described from the Santa Monica Mountains on the slope below the Hollywood sign (Hasse 1913). Rare (F). Lecidea fuscoatra (L.) Ach. - Saxicolous, terricolous. Common (F & W). Lecidea laboriosa Mttll. Arg. - Saxicolous. Common (F & W). Lecidea tessettata Florke - Saxicolous. Common montane species above 6000 feet. Rare in survey area (F). Lecidella as etna (Nyl.) Knoph & Hertel - Saxicolous, rarely terricolous. Common (F & W). 74 Crossosoma 35(2), Fall -Winter 2009 Lecidella stigmalea (Ach.) Hertel & Leuckert - Saxicolous. Infrequent (F). Lepraria adhaerens K. Knudsen, Elix & Lendemer- Saxicolous, terricolous. Common (F & W). Lepraria borealis Lohtander and Tonsberg - Saxicolous. Rare on moss and sandstone. (W). Lepraria caesioalba (B. de Lesd.) J.R. Laundon - Coticolous, saxicolous, terricolous. Two chemotypes. Frequent (F & W). Lepraria santamonicae K. Knudsen & Elix - Saxicolous, terricolous. Frequent (F & W). Lepraria texta K. Knudsen, Elix & Lendemer - Saxicolous, terricolous. Frequent (F), Lepraria xeropltila Tonsberg - Terricolous. Common maritime species. Rare (W). Leptogium arsenei Sierk - Saxicolous. Rare (W). Leptogium palmatum (Huds.) Mont. - Terricolous. Rare (W). Leptogium plicatile (Ach.) Leight. - Saxicolous. Single small population on wet shaded calcareous sandstone in oak woodland. Rare (W). Leptogium tenuissimum (Dicks.) Korb. - Terricolous. Rare (W). Lichenoconium lecanorae (Jaap) D. Hawksw. - Lichenicolous fungus on apothecia of Lecanora muralis. Frequent (F). Lichenoconium lichenicola (P. Karst.) Petrak & H. Sydow - Lichenicolous fungus infrequent on Physcia dimidiata (F). Licit enostigma cosmopolites Hafellner & Calat. - Lichenicolous fungus frequent on Xanthoparmelia species (F). Licit enostigma rugosum Thor - Lichenicolous fungus frequent on Diploschistes species (F). Licit enostigma subradians Hafellner, Calat. & Nav.-Ros. - Lichenicolous fungus frequent on Acarospora social is (F). Micarea microccoca (Korb.) Gams ex Coppins - Corticolous. Rare on mature bark of old half-dead Sambucus mexicana (F). Miriquidica scotopholis (Tuck.) B.D.Ryan & Timdal - Saxicolous. Common (F & W). Peltula bolanderi (Tuck.) Wetmore - Saxicolous. Rare (W). Peltula euploca (Ach.) Poelt - Saxicolous. Infrequent (F). Peltula obscurans var. hassei (Zahlbr.) Wetmore - Saxicolous. Infrequent (F). Peltula obscurans (Nyl.) Gyeln. var. obscurans - Saxicolous. Rare (W). Phaeophyscia hirsuta (Mereschk.) Essl. - Corticolous. Infrequent on Ouercus agrifolia (W). Phaeophyscia sciastra (Ach.) Moberg - Saxicolous. Rare (W). Physcia adscendens (Fr.) H. Olivier - Corticolous, infrequently saxicolous. Common on Quercus agrifolia , chaparral and coastal sage shrubs (F & W). Crossosonui 35(2), Fail -Winter 2009 75 Physcia biziana (A. MassaJ) Zahlbr. - Corticolous. Rare on Adenostoma fasciculatum (W). Physcia dimidiata (Arnold) Nyl. - Saxicolous. Common (F & W). Physcia iribacia (Ach.) Nyl. - Saxicolous. Infrequent (F & W). Physconia isidiigera (Zahlbr.) Essl. - Corticolous, saxicolous. Frequent on sandstone and Quercus agrifolia (F & W). P/acidium lacinulatum (Ach.) Breuss - Terricolous. Common (F & W). Placidiunt squamulosum (Ach.) Bruess - Terricolous. Infrequent (W). Placopyrenium noxium Breuss - Saxicolous. Second known population in California (Breuss 2009). Rare (F). Placynthiella hyporhoda (Th. Fr.) Coppins & R James - Terricolous. Infrequent (W). Placynthiella uliginosa (Schrad.) Coppins & P. James - Terricolous, saxicolous. Frequent (W). Polysporina simplex (Davies) Vllzda - Saxicolous. Common (F & W). Polysporina subfucescens (Nyl.) K. Knudsen & Kocourk. -Lichenicolous fungus common on saxicolous crustose lichens (F & W). Psora californica Timdal - Terricolous. Infrequent (F & W). Psora luridella (Tuck.) Fink - Saxicolous, Terricolous. Frequent (F & W). Psorothicia montinii (A. Massal.) Forss. - Saxicolous. Infrequent (W) Psorotichia schaereri (A. Massal.) Arnold - Terricolous. Infrequent (W). Rinodina species - Terricolous. Endemic to California and will be described by John Sheard. Rare (F & W). Rinodina gennarii Bagl. - Saxicolous. Maritime species, infrequent on concrete of old flood control wall and on rhyolite (F & W). Rinodina intermedia Bagl. - Terricolous. Infrequent (F & W). Sarcogvne arenosa (Herre) K. Knudsen & Standley - Saxicolous. Common (F&W). Sarcogyne plicata H. Magn. - Saxicolous. The name Sarcogyme privigna (Ach.) A. Massal. was applied to this taxon (Knudsen & Standley 2007) and S. plicata , described from Upland, was treated as a synonym (Knudsen & Lendemer 2005). Recent unpublished molecular analysis of specimen from Fremont Canyon showed it was not conspecific with S. privigna in Fennoscandia (Westberg, pers. comm.) In this paper we resurrect S. plicata, but it may not apply to all taxa in California. Frequent (F & W). Sarcogyne reebiae K. Knudsen - Saxicolous. Described from Weir Canyon (Knudsen & Standley 2007). Infrequent ( W). Sarcogyne regularis Korb. - Saxicolous. Infrequent (W). Sarcogyne simi/is H. Magn. - Saxicolous. Common (F & W). Strangospora deplanata (Almq.) Clauz. & Cl. Roux - Corticolous. Rare on mature bark of old half-dead Sambucus mexicana (F). 76 Crossosoma 35(2), Fall -Winter 2009 Stigmidium fuscatae (Arnold) R. Sant. - Lichenicolous fungus on A car os par a obpallem. We will revise this taxon in southern California. Rare (F). Stigmidium squamariae (B. de Lesd.) Cl. Roux & Triebel- Lichenicolous fungus on apothecia of Lecanora muralis. Infrequent (F). Sligmidium xanthoparmelium Hafellner - Lichenicolous fungus on Xanthopamelia species. Infrequent (F). Teloschisles chrysophthalmus (L.) Th. Fr. -Corticolous. Rare on Malacothamnus fasciculatus (W). Thelomma mammosum (Hepp ex Hartung) Tibell - Saxicolous. Coastal species. Infrequent (W). Thelomma santessonii Tibell - Saxicolous. Maritime species. A single population was discovered on ridge of Fremont Canyon (F). Toninia trislis (Th. Fr.) Th. Fr. - Saxicolous. Infrequent. (F). Trapelia coarctata (Turner ex Sm.) M. Choisy- Saxicolous, terricolous. Frequent (F & W). Trapelia glebulosa (Sw.) J.R. Laundon - Saxicolous, terricolous. Frequent (F & W). Trapelia placodioides Coppins & P. James - Saxicolous, terricolous. Locally common, but probably rare in survey area after the Weir Canyon fire because it occurred on slope within dense chaparral (W). Trapeliopsis hisorediata McCune & Camacho - Saxicolous, terricolous. Rare on sandstone (W). Trapeliopsis flexuosa (Fr.) Coppins & P. James - Lignicolous. On dead and old burnt wood (F & W). Trapeliopsis glaucopholis (Nyl. ex Hasse) Printzen & McCune - Saxicolous, terricolous. Frequent (F & W). Trapeliopsis steppica McCune & Camacho - Saxicolous, terricolous. This is the second population discovered in Santa Ana Mountains. Rare (F). Umhilicaria phaea Tuck - Saxicolous. Infrequent. (F). Verrucaria calkinsiana Servit - Saxicolous. Common (F & W). Verrucaria furfuracea (B. de Lesd.) Breuss. - Saxicolous. Frequent (W). Verrucaria fusca Pers. ex Ach. - Saxicolous. Frequent (F & W). Verrucaria fuscoatroides Servit - Saxicolous. Infrequent (W). Verrucaria memnonia (Flot. ex Korb.) Arnold - Saxicolous. On small hard granite rocks washed out of sandstone deposits. Infrequent (F & W). Verrucaria onegensis Vain. - Saxicolous. Rare (F). Verrucaria turgida Servit - Saxicolous. Rare (F). Verrucaria viridula (Schrader) Ach. - Saxicolous. Infrequent (F). Xanthoparmelia amahleana (Gyeln.) Hale - Saxicolous. Infrequent (F). Xauth oparmelia cumherlandi (Gyeln.) Hale - Saxicolous. Common (F & W). Xanthoparmelia lineola (E.C. Berry) Hale - Saxicolous. Infrequent (F). Crossosoma 35(2), Fall-Winter 2009 77 Xanth oparmelia mexicana (Gyeln.) Hale - Saxicolous. Common (F & W). Xanthoparmelia neotartica Hale - Terricolous. Infrequent (F & W). Xanth oparmelia novomexicana (Gyeln.) Hale - Saxicolous. Infrequent (F). Xanthoparmelia suhplitti Hale - Saxicolous. Infrequent (F). Xanthoparmelia verruculifera (Nyl.) O. Blanco, A. Crespo, Elix, D. Hawksw. & Lumbsch - Saxicolous. Rare (F). Xanthoria polycarpa (Hoffm.) Th. Fr. ex Rieber - Corticolous. Infrequent on Malacothamnus fasciculatus and Adenostoma fasciculatum (W). Xanthoria ten ax L. Lindblom - Corticolous. Common on willows and chaparral before the Weir Canyon fire (W). CONCLUSIONS We report 169 taxa in 56 genera for the northwestern Santa Ana Mountains, comprising 156 lichens and 13 lichenicolous fungi. Two species were described new to science, Sarcogyne reebiae from Weir Canyon (Knudsen & Standley 2007) and Gelatinopsis acarosporicola from Fremont Canyon (Kocourkova & Knudsen 2009). Four recently described species have paratype locations in Fremont Canyon and Weir Canyon: Lepraria adhaerens (Knudsen et al. 2007), L. santaemonicae (Knudsen & Elix 2007), L. texta (Knudsen & Elix 2007), and Leccmora mimzii (Knudsen & Lendemer 2009). John W. Sheard (SASK) is currently in the process of describing the terricolous Rinodina species included in the checklist which is also known from the Santa Monica Mountains. Previously Carbonea latypizodes (Knudsen et al. 2008), Cercidospora caudata (Etayo et al. 2007). Lichenostigma rugosnm (Knudsen & Kocourkova 2008) and Trapelia plcicodioides (Knudsen 2006) were reported new for California from Fremont Canyon and Weir Canyon. We believe the total species diversity would have been higher if Fremont Canyon had been surveyed before the devastating fire. Some lichens considered rare in Weir Canyon may have been locally extirpated by the fire of 2007, especially species on bark and wood. While the authors consider plants and animals the most important biological resources on public lands, lichens are a significant resource in relatively undisturbed habitats. Over 1500 taxa of lichens, lichenicolous fungi, and allied microfungi have been reported from California (Tucker & Ryan 2006) and the numbers of known taxa increase regularly, despite any reductions of numbers through taxonomic advances. For instance, based on continuing research, 298 taxa occur in the Santa Monica Mountains (Knudsen & Kocourkova 2009) and 387 taxa on Santa Rosa Island (Knudsen unpublished). The lichens and lichenicolous fungi we report in this paper may comprise fewer than half of the species that occur in the Santa Ana Mountains. We expect the total diversity of lichens and 78 Crossosomii 35(2), Fall-Winter 2009 lichenicolous fungi to be at least 280 species, over one fourth of the total diversity of the vascular flora (which includes native and non-native plants). All lichens and lichenicolous fungi reported here are apparently native to the range. Only onefloristic study of lichens has been published on the Santa Ana Mountains, based on a short survey of the Santa Rosa Plateau (Weber & Bratt 1987). Many of that paper’s taxonomic concepts and species concepts are now archaic due to rapid advances in lichen taxonomy spearheaded by the Sonoran lichen flora project (Nash et al. 2002, 2004, 2007). Weber and Bratt reported approximately 78 species. Without a revision of the specimens and based on our knowledge of the Santa Rosa Plateau, approximately 49 species they reported also occur in the northwestern Santa Ana Mountains. The main difference is the lichen flora on oaks is more diverse on the Santa Rosa Plateau because of apparently higher relative annual humidity (particularly from fog incidents) and the occurrence of Engelmann oaks and scrub oaks besides Quercus agrifolia. Our checklist can be used as the foundation of a preliminary checklist of the whole range as well as for Orange County. This paper represents the first installment of our continuing floristic study of the Santa Ana Mountains. Our next paper will be a study of the Santa Rosa Plateau. Before grazing and the dominance of invasive plants, the valley grasslands in Fremont Canyon and Weir Canyon may have supported more extensive biological soil crusts of lichens and bryophytes, native grasses and annuals. Terricolous lichens were probably common in biological soil crusts and included species extirpated from study area like Aspicilia praecrenata (Nyl. ex Hasse) Flue. Most terricolous lichens are now rare or infrequent. Terricolous lichens now subsist as relics in microhabitats on thin-soiled slopes, on decaying sandstone, or on consolidated soil in pockets on sandstone outcrops. The most common terricolous lichen we found in Weir and Fremont Canyons was the pioneering Cladonia nashii, which can grow on slopes of loose alluvium and road cuts as well on decaying or re-consolidated sandstone. The Tecate cypress (Cupressus forebesii) grove was visited before the Fremont Canyon fire devastated it. The trees did not support lichens. Two lichens, Lecanora munzii and L. simeonensis , were collected in upper Weir Canyon. These species are lignicolous, growing on the wood of older chaparral and coastal sage shrubs. They are rare, like Cyphelium brachysporum Nadv. (Lendemer et al. 2008) described from Murrieta, probably because more frequent fires are reducing mature stands where their substrates are found. Both these species may have been extirpated from Weir Canyon by the fire of 2007 and were Crossosonui 35(2), Fall -Winter 2009 79 not documented from Fremont Canyon. A number of maritime species, common on the coast of southern California and on the Channel Islands, occur in the northwestern Santa Ana Mountains, approximately 14-17 miles (22-27 km) inland from the coast. All were rare or infrequent. We only report 13 species of lichenicolous fungi. The number of lichenicolous fungi reported here does not include at least five taxa probably new to science we discovered in Fremont Canyon (after the fire) which were all rare. We were unable to collect sufficient material for descriptions. The first, Endococuus thelommae Kocourk. & K. Knudsen, has been accepted for publication in Germany in a forthcoming volume of Bibliotheca Lichenologica. Sufficient type material was finally collected on Santa Rosa Island. Lichens grow very slowly, especially in Mediterranean climates based on our observations, with most lichens growing only micro-millimeters in an average rain year and probably not at all during drought episodes. The recovery of the lichen biota in the northwestern Santa Ana Mountains will be a slow process taking decades. It is tragic that both of these fires were caused by humans, the Weir Canyon fire by arson and the Fremont Canyon fire through mismanagement of a controlled bum. ACKNOWLEDGMENTS We thank our reviewers, Brendan Hodkinson (DUKE) and Caleb Morse (KANU). We thank for their help with identifications Othmar Breuss (Austria), Jack Elix (Australia), J.C. Lendemer (NY), Christian Printzen (Germany) and John Sheard (Canada). We thank ecologist Trish Smith (Nature Conservancy) for her constant support of our research. This project was supported by grants from The Nature Conservancy. LITERATURE CITED Arup, U. 2009. The Caloplaca holocarpa group in the Nordic countries, except Iceland. Lichenolagist 41: 111-130. Boyd, S., T.S. Ross and F. M. Roberts, Jr. 1995. Additions to the vascular flora of the Santa Ana Mountains, California. Alisa 14(2): 105-108. Boyd, S., T.S. Ross and D. Bramlett. 1995. Vascular flora of the San Mateo Wilderness Area, Cleveland National Forest, California. Alisa 14(2): 109-139. 80 Crossosoma 35(2), Fall -Winter 2009 Breuss, O. 2009. A synopsis of the lichen genus Placopyrenium (Verrucariaceae), with descriptions of new taxa and a key to all species. In: Biodiversity and ecology of lichens — Liber Amicorum Harrie Sip man. Aptroot, A., M. R. D. Seaward and L. B. Sparrius (eds.) Bibliotheca Lichenologica 99: 93-112. J. Cramer, Berlin & Stuttgart, 2009. Etayo, J., J. Kocourkova and K. Knudsen. 2007. New records of lichenicolous fungi for California. Bulletin of the California Lichen Society 14(2): 37-39. Hasse, H. E. 1913. The lichen flora of southern California. Contributions from the United States National Herbarium 17: 1-132. Knudsen, K. 2006. Noteworthy collections: Lichens and lichenicolous fungi. Crossosoma 32( 1 ): 31-33. Knudsen, K. 2007. Acarospora. In: T. H. Nash, III, C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3, pp. I -38. Lichens Unlimited. Arizona State University, Tempe. Knudsen, K. and J. A. Elix. 2007. A new Lepraria (Stereocaulaceae) from the Santa Monica Mountains in southern California. The B/yologist 1 10(1): 115-118. K. Knudsen and J. A. Elix. 2007. Lepraria. In: T. H. Nash, III, C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3. pp. 384-390, Lichens Unlimited, Arizona State University, Tempe. Knudsen, K., J. A. Elix. and J.C. Lendemer. 2007. Lepraria adherens : a new species from North America. Opuscula Philolichenum 4: 5-10. Knudsen, K. and J. Kocourkova. 2008. New records of lichen and lichenicolous fungi from California. Crossosoma 34(1): 37-39. Knudsen, K. and J. Kocourkova. 2009. Lichens, Lichenicolous and Allied Fungi of the Santa Monica Mountains, Part 4: Additions and Corrections to the Annotated Checklist. Opuscula Philolichenum 7: 29-48. Knudsen. K. and J. C. Lendemer. 2005. Changes and additions to the checklist of North American lichens - III. Mycofaxon 93: 277-28 1 . Knudsen, K & J. C. Lendemer. 2009. Two new species of Lecanora with gyrophoric acid from North America. Opuscula Philolichenum 7: 21-28. Knudsen, K., B. Owe-Larsson, J. A. Elix, J. C. Lendemer and J. Kocourkova. 2008. Lichens and lichenicolous fungi of the Santa Monica Mountains, Part 3: additions and corrections to the annotated checklist. Opuscula Philolichenum 5: 53-60 Crossosoma 35(2), Fall-Winter 2009 81 Knudsen, K., V. Reeb, M. Westberg, R. Srikanthaand D. Bhattacharya. 2010. Acarospora rosulata in Europe, North America and Asia. Lichenologist 42(4): not assigned. Knudsen, K. and S. M. Standley. 2007. Sarcogyme. In: T. H. Nash, III. C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3, pp. 289-296. Lichens Unlimited, Arizona State University, Tempe. Kocourkova, J. and K. Knudsen. 2009. Gelatinopsis acaarosporicola (Helotiaceae), a new lichen icolous fungus on Acarospora social is from western North America. The Biologist 112(2): 363-367. Lathrop, E. W. and R. F. Thome. 1978. A Flora of the Santa Ana Mountains, California. Aliso 9(2): 197-278. Lendemer, J.C. and K. Knudsen. 2009. Two new usnic acid-containing species of Lecanora from western North America. Opuscula Philolichenum 6: 73-80. Lendemer, J.C., J. Kocourkova and K. Knudsen. 2008. Studies in lichens and lichenicolous fungi: notes on some taxa from North America. Mvcotaxon 105: 379-386. Nash III, T.H., B.D. Ryan, C. Gries, and F. Bungartz (eds.) 2002. Lichen Flora of the Greater Sonoran Desert Region. Vol. I, Tempe, Arizona: Lichens Unlimited, Arizona State University, 532 pp. Nash III, T.H., B.D. Ryan, P. Diederich, C. Gries, and F. Bungartz (eds.) 2004. Lichen Flora of the Greater Sonoran Desert Region, Vol. 2, Tempe, Arizona: Lichens Unlimited, Arizona State University, 744 pp. Nash III, T.H., C. Gries, and F. Bungartz (eds.) 2007 (2008). Lichen Flora of the Greater Sonoran Desert Region, Vol. 3, Tempe, Arizona: Lichens Unlimited, Arizona State University, 567 pp. Tucker, S.C. and B.D. Ryan. 2006. Constancea 84: Revised Catalog of Lichens, Lichenicoles. and Allied Fungi in California (http://ucjeps.berkeley. edu/ constancea/84/). Acccessed May, 2009. Weber, W. A. and C. Bratt. 1987. Lichens and bryophytes of the Santa Rosa Plateau Nature Conservancy Reserve, Riverside County, California. Evans ia 4: 21-25 Wetmore, C. M. 2007. Caloplaca. In: T. FI. Nash, III, C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3, pp. 179-220. Lichens Unlimited, Arizona State University, Tempe. 82 Crossosoiiui 35(2), Fall -Winter 2009 NEW RECORDS OF LICHENS AND LICHEN1COLOUS FUNGI FOR CALIFORNIA I. Jana Kocourkova University of Life Sciences, Faculty of Environmental Sciences, Department of Ecology Kamycka 129 165 21 Praha 6 - Suchdol, Czech Republic e-mail: kocourkovaj@fzp.czu.cz Kerry Knudsen The Herbarium, Dept, of Botany & Plant Sciences, University of California, Riverside, California 92521 kk999@msn.com Othmar Breuss Naturhistorisches Museum, Botanische Abteilung, Burgring 7, A-1010 Wien, Austria obreuss@bg9.at ABSTRACT: Three species of lichens, two which occur in intertidal zone, and three lichenicolous fungi are reported new for California: Arthonia epiphyscia, Dactvlospora saxatilis , Rhizocarpon effiguratum, Unguiculariopsis lettaui, Verrucaria prominula, and Wahlenbergiel/a striatula. KEYWORDS: Biodiversity, Co/lemopsidiutn, rising sea levels. INTRODUCTION In 2008 we began studying the intertidal zone for lichens in southern and central California with Othmar Breuss of Austria. We are concerned with what species occur in California and the impact of rising sea levels on their diversity. So far lichens are rare in the intertidal zone in southern and central California, with algae winning the competition for intertidal space. The cyanolichen Collemopsidium sublittove (Leighton) Grube & B. D. Ryan is the most common species. In this paper we report two new records of intertidal lichens for California. Lichenicolous fungi are a successful and diverse group of fungi which are growing on the thallus and apothecia of lichens. Generally they occur in habitats that have long ecological continuity. Over 1500 species of lichenicolous fungi species are known to science (Lawrey and Diederich 2003). In 2008 we recognized 108 described species of lichenicolous fungi as occurring in California (Kocourkova Crossosoma 35(2), Fall -Winter 2009 83 & Knudsen, unpublished), excluding lichenicolous lichens, some allied fungi and dubious reports that we have not verified (Tucker & Ryan 2006). In publications of this year, including this paper, we reported 13 species new to California of which 6 were new to North America (Lendemer et al. 2009; Kocourkova 2009; Kocourkova & Knudsen 2009c; Knudsen & Kocourkova 2009) and described 5 new species which occur in California (Knudsen et al. 2009; Knudsen & Kocourkova 2009; Kocourkova 2009; Kocourkova and Knudsen 2009a & 2009d) for a total of 127 species. We expect the reported diversity of lichenicolous fungus species for the state to at least double in next ten years. Taxa new to science are still easily found in California and we are in the process of describing over a dozen species from the state, with three already in review or press. Our good friend and colleague Paul Diederich of Luxembourg will probably describe at least three new species from central California. The Species 1. Arthonia epiphyscia Nyl. has a widespread distribution and is known from Africa, Asia, Europe, and North and South America, and is restricted to species of Pty’scia (Kocourkova 2000; Grube 2007). It has distinctive superficial black ascomata that are easily seen on the white thallus of Physcia species. Though the host Physcia adscendens is one of the most common lichens in California on coastal sage shrubs and chaparral as well as oaks, A. epiphyscia is proving to be rare. The collection was made during the Bioblitz held by the San Diego Natural History Museum in 2009. Specimen examined: San Diego Co.: San Diego, Mission Trails Regional Park, Visitor Center Loop Trail, 32° 49’ 18’' N, 117° 03’ 18” W, 84 in, on Physcia adscendens on Artemisia californica , May I, 2009, Knudsen 10970 (UCR). 2. Dactylospora saxatilis (Schaerer) Hafellner is a lichenicolous fungus widespread on Pertusaria species in Europe, North America, Asia and Africa (Hafellner 1979, 2004). It has black lecideine apothecia with dark brown one-septate ascospores and asci with an external gelatinous sheath which is 1+ blue (euamyloid). It was rare on Pertusaria on Santa Rosa Island, but Skyttea pertusariicola Diederich & Etayo was frequent. Specimen examined: Santa Barbara County: Santa Rosa Island, Channel Islands National Park, hillside above Beecher’s Bay, 33° 59’ 13” N, 120° 01’ 13” W, 46 m, on Pertusaria Jiavicunda, Oct. 15, 2006, Knudsen 7496.2 (UCR). 84 Crossosoma 35(2), Fall -Winter 2009 3. Rhizocarpon effiguratum (Anzi) Th. Fr. is a lichen which is often parasitic on P leopsidium flavum (Bellardi) Korb. (which is a yellow lichen common on granite in southern California above 6000 feet). Previous reports from California were questionable (Tucker & Ryan 2006) and it is currently known from southwestern North America only from Arizona in the San Francisco Peaks and Mt. Baldy in the White Mountains (Feuerer & Timdal 2007). Rhizocarpon effiguratum was rare on the top of Santa Rosa Mountain. Specimen examined: Riverside County: Santa Rosa Mountain, ridge toward Toro Peak, 33° 32’ 11” N, 116° 27’ 24” W, 2416 m, Jul. 18, 2009 Knudsen 11499 w / Tom Chester et al. (UCR, MIC). 4. Unguiculariopsis iettaui (Grummann) Coppins is a parasymbiotic or slightly parasitic fungus often forming galls on the fruticose lichen Evernia prunastri. It is wide-spread in Europe and Macronesia (Diederich & Etayo 2000). It was recently reported new from North America from a single collection from Oregon (Diederich 2002). We report the second collection from North America from Marin County. The black ascomata have distinct appendages and simple hyaline ascospores 6-8 x 3-4 pm. Specimen examined: Marin County: Point Reyes National Seashore; Earthquake Trail across road from the Bear Valley Visitors’ Center, near wood bridge over brook, 38° 04’ N 122° 79’ W, 3 1 m, on Evernia prunastri on bark of Alnus rubra , July 10, 2008, Kocourkovd & Knudsen, IAL Excursion (PRM 915120). 5. Verrucariu prominula Nyl. was collected on Point Loma in the upper littoral zone on sandstone, where it is rarely submerged. It is distinguished by its relative small ascospores and relatively large perithecia with thick involucrellum and is known from Asia, Europe, North America and Tasmania (Orange et al. 2009). It is new to California. Specimen examined: San Diego County: Point Loma, Cabrillo National Monument: tidepools, 32° 40' 8” N 117° 14' 42” W, 0.5 m, July 30, 2008, Knudsen 10685 w/ Kocourkovd (UCR). 6. Wahlenbergiella striatula (Wahlenb.) Gueidan & Thus occurred in the intertidal zone with red algae, green seaweed, and barnacles on basalt on the western tip of West Anacapa below Rat Rock and has a subgelatinious greenish-black thallus with black spots or ridges. Though cosmopolitan (Orange et al 2009), it is new for California. Crossosonui 35(2), Fall-Winter 2009 85 Specimen examined: Ventura County: West Anacapa Island, Channel Islands National Park, below Rat Rock 34° O’ 50” N 1 19° 26' 32” W, 0 m, on basalt, Nov. 19 2008, Knudsen 10685 w/ Kocourkova (UCR). ACKNOWLEDGEMENTS We thank Sarah Chaney (NPS botanist) for guiding us on our exploration of West Anacapa Island. The work of Jana Kocourkova was supported by the University' of Life Sciences, Faculty of Environmental Sciences, Prague, Czech Republic. The work of Kerry Knudsen was supported in part by a co-operative agreement between Channel Islands National Park and the University of California at Riverside. LITERATURE CITED Diederich, P. 2002. New species and new records of American lichenicolous fungi. Herzogia 16: 41—90. Diederich, P. and J. Etayo. 2000. A synopsis of the genera Skyttea, Llimoniella , and Rhymbocarpns (lichenicolous Ascomycota, Leotiales). Lichenologist 32(5): 423-485. Grube, M. 2007. Arthonia. In: T. H. Nash, III, C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3, pp. 39- 61, Lichens Unlimited, Arizona State University, Tempe. Feuerer, T. & E. Timdal. 2004. Rhizocarpon. In: T. H. Nash, III, C. Gries and F. Bungartz (eds.) Lichen Flora of the Greater Sonoran Desert Region. Volume 3, pp. 456-466, Lichens Unlimited, Arizona State University7, Tempe. Hafellner, J. 1979. Karschia. Revision einer Sammelgattung an der Grenze von lichenisierten und nichtlichenisierten Ascomyceten. Nova Hedwigia, Beih. 62: 1-248. Hafellner J. 2004. Dactvlospora. In: T.H. Nash, III., B. D. Ryan, P. Diederich, C. Gries, and F. Bungartz, F. (eds.), Lichen Flora of the Greater Sonoran Desert Region, Vol. 2, pp. 645-648. Lichens Unlimited, Arizona State University, Tempe, Arizona. Knudsen, K. and J. Kocourkova. 2009. Lichens, Lichenicolous and Allied Fungi of the Santa Monica Mountains, Part 4: Additions and Corrections to the Annotated Checklist. Opuscula Philolichenum 7: 29-48. Knudsen, K., Kocourkova, J. and J. Etayo. 2009. A new species of Sphaerellothecium (Mycosphaerellaceae) on Placidium lacinulatum. Opuscula Philolichenum 6: 4 1 — 44. 86 Crossosomii 35(2), Fall-Winter 2009 Kocourkova, J. 2000. Lichenicolous fungi of the Czech Republic (The first commented checklist). Shornik Ndrodnlho Musea v Praze, Rada B 55(3- 4): 59-169. Kocourkova, J. 2009. Observations on the genus Neolamya, with the description of the new species N. xanthoparmeliae (Ascomycota, genera incertae sedis). Opus cula PhUolichenum 6: 137—148. Kocourkova, J. and K. Knudsen 2009a. Gelatinopsis acarosporicola (Helot'mceae), a new lichenicolous fungus on Acarospora socialis from western North America. The Bryologist 112(2): 363—367. Kocourkova, J. and K. Knudsen. 2009b. Stigmidiwn epistigmellum (Mycosphaerellaceae), a lichenicolous fungus from maritime Caloplaca in North America. The Bryologist 112(3): 578-583. Kocourkova, J. and K. Knudsen. 2009c. Three Lichenicolous Fungi new for North America. Evamia 26(3): 1 48- 151. Kocourkova, J. and K. Knudsen. 2009d. A new species of Stigmidiwn (Mycosphaerellaceae, Ascomycetes) from western North America. Czech My colog}' 61(1): 73-80. Lawrey, J. D. and P. Diederich. 2003. Lichenicolous fungi: interactions, evolution, and biodiversity. The Bryologist 106: 81-120. Lendemer, J. C., J. Kocourkova and K. Knudsen. 2008: Studies in lichens and lichenicolous fungi: more notes on some taxa from North America. My cot axon 108: 491-497. Orange, A., D. L. Hawksworth, P. M. McCarthy and A. Fletcher. 2009. Verrucaria. In C.W. Smith, A. Aptroot, B. J. Coppins, A. Fletcher, O. L. Gilbert, P.W. James and P. A. Wolseley (eds.) The Lichens of Great Britain and Ireland, pp. 931-957, Slough, United Kingdom: Richmond Publishing Company. Tucker, S. C. and B. D. Ryan. 2006. Revised catalog of lichens, lichenicoles, and allied fungi in California. Constancea 84: 1-275 + 1-52. Crossosoma 35(2), Fall -Winter 2009 87 COST AND EFFECTIVENESS OF SMALL-SCALE FOENICULUM VULGARE CONTROL METHODS Abigail Gwinn 2 Walker Ct. Soledad, CA 93960 abigail_gwinn@csumb.edu ABSTRACT Information on effective methods of controlling invasive plants is crucial to land managers and others who seek to restore their lands to a more natural state. Much of the literature on invasive plant removal techniques focuses on large invasions and explores the methods appropriate for those situations. However, information regarding small invasions and effective control methods utilizing the limited materials available to individuals and small organizations is less widely available and often anecdotal. This study, conducted during the summer of 2006 at two sites in Salinas, California, examined three methods of controlling a small- scale infestation of fennel ( Foemculum vulgare) to identify which would be most effective and economical. The methods were: digging out individual plants with a shovel, chopping the plants repeatedly with a machete, and chopping the plants and immediately spraying the stumps with the herbicide Rodeo. Digging up each plant was the most effective method; chopping and spraying was a close second. Chopping repeatedly without herbicide treatment was not effective, resulting in apparent mortality7 rate very similar to that seen among untreated plants. Of the two effective methods, chopping and spraying was the least expensive. Digging the individual plants was the most time consuming and the most expensive. My recommendation for controlling F. vulgare based upon these results is to chop and spray when soils are hard and herbicide application is permitted, and to dig up each plant when soils are soft or where herbicide use is not an option. KEYWORDS Foeniealum vulgare, fennel, anise, small-scale invasions, glyphosate, Rodeo, herbicide, invasive plants, cost-effectiveness INTRODUCTION Invasive plants are defined as non-native plants that have been introduced to an area and are able to produce reproductive offspring at considerable distance from the parent plant, creating the potential to spread extensively (Richardson et al. 2000). Estimated economic costs associated with invasive species in the United States include $97 billion in damage to agriculture, forestry, and the environment by 79 species between 1906 to 1991 (Pimental et al. 2000) and $ 125 billion spent 88 Crossosoma 35(2), Fall-Winter 2009 on control per year (Myers and Bazely 2003). Though some plant expansion beyond native environments occurs naturally due to climate and habitat changes, invasions have been caused by intentional or unintentional introduction by human activity (Luken and Thieret 1997; Pysek et al. 2004). Invasive plants are estimated to occupy over 100 million acres in the United States and to be spreading at a rate of 3 million acres per year (Myers and Bazely 2003). Foemculum vulgare Miller, commonly known as fennel, sweet fennel, aniseed, anise or sweet anise, was introduced to California from the Mediterranean region at least 120 years ago (Klinger 2000). It generally occurs in open, disturbed areas or along roadsides (Brenton and Klinger 2002). Foemculum vulgare has some value as an agricultural crop (Brenton and Klinger 2002) and has likely escaped from cultivation several times (Klinger 2000). The California Invasive Plant Council ranks F. vulgare high on the California Invasive Plant Inventory Database, reflecting the species’ level of adverse ecological impact in California (California Invasive Plant Council 2009). Foemculum vulgare is a perennial herb that grows rapidly and spreads by seed or re-growth from its root crown (Klinger 2000). A single F. vulgare plant can produce thousands of seeds during its first growing season, and hundreds of thousands of seeds during the plant’s second year (Erskine Ogden and Rejmanek 2005). Foemculum vulgare may compete with other plants for resources (Holloran et al. 2004) or exclude them altogether by creating nearly monotypic stands (Bell et al. 2008). It successfully competes with native perennials in coastal sage communities, probably due in part to its ability to grow during the summer when most native species are dormant (Weber 2003). The ability of F. vulgare to re-sprout from its root stock can hinder determination of plant mortality because seemingly dead plants may resprout the following year, as illustrated in Bell et al. (2008). The purpose of this project was to test three F. vulgare control methods to determine which worked most effectively and economically at two locations in Salinas, California: Natividad Creek Park and Upper Carr Lake. Klinger (2000) recommended controlling light infestations by digging out individual plants. However, in heavy clay soils, as found at both sites, digging up each plant is labor intensive. Several studies of large-scale F. vulgare control on Santa Cruz Island, California, have included controlled burns and aerial herbicide application (Brenton and Klinger 2002; Erskine Ogden and Rejmanek 2005), neither of which is appropriate for small-scale infestations in public parks within city limits. Herbicides that have been effective in controlling F. vulgare are: amine and ester formulations of triclopyr (Garlon 3A and Gallon 4), and glyphosate as Roundup (Klinger 2000). Crossosoma 35(2), Fall-Winter 2009 89 I partnered with Return of the Natives (RON) to use mo of their restoration areas as study sites. Return of the Natives also provided the tools and materials used in this project. Return of the Natives is a non-profit restoration and education project that has been working to restore native habitats by removing invasive plants and planting natives at Natividad Creek Park since 1995 and Upper Carr Lake since 2003. METHODS Study Area Upper Carr Lake includes a large pond frequented by water birds, a bike path, and several restored areas, including the hillside near a county yard. This hillside is the location of one Foemculum vulgare invasion in the park and served as one study site. Natividad Creek Park contains common city park facilities as well as several open fields with a mix of native and exotic plant life. One of these fields and an adjacent hillside, bordered by Freedom Boulevard and Nogal Drive, is another F. vulgare location, selected as my second study site. Each site was approximately one acre in area, with widely spaced, typically young F. vulgare individuals spread throughout. Clumps of F. vulgare tended to be small, with the plants widely dispersed. When measured on 26 June 2006, most specimens were taller than 120 cm. Plant circumference was measured at 20 cm above the ground; most plants in the study had a circumference of 10 to 40 cm. The soil at both sites was very hard-packed, and the surrounding vegetation consisted of non-native annual grasses with a few native bunch-grasses and young oak trees ( Quercus sp.), coyote brush ( Baccharis pilularis), and other native trees, shrubs, and grasses that were planted as part of a restoration program for the park. Other invasives, such as curly dock (Rum ex crispus ), bristly oxtongue ( Picris echioides ), and English plantain ( Plantago lanceolata ), were also present. Experimental Design I randomly selected 50 Foeniculum vulgare plants at Natividad Creek Park and 50 plants at Upper Carr Lake. Each selected plant was randomly assigned to one of four different treatment methods, with 25 plants per method. Treatment began on 10 July 2006. The methods were; • Control: No treatment other than removal of any flower heads to prevent seeding. • Chop Repeatedly: Plants were chopped with a machete to 20 cm or less above the ground. They were revisited every' two weeks to chop any new growth. 90 Crossosonia 35(2), Fall -Winter 2009 • Chop and Sprav: Plants were chopped with a machete to 20 cm or less and then immediately sprayed with the herbicide Rodeo. This was the only treatment method with two people working together on the same plant. These plants were not retreated. • Dig: Plants were dug up with a shovel to remove as much of the root and rootcrown as possible. Time spent on each plant for all methods except the control was recorded to calculate the cost of labor (Results section, below). For the chop and spray treatment I used 25 ml of Rodeo mixed with water and no surfactant in a standard, one-liter spray bottle. Half of the mixture remained after treating all 25 plants in the treatment group. Treatment was halted on 4 September 2006, after a final round of chopping and an assessment of all plants. The plants were visually evaluated and placed into one of three categories: • Apparent mortality: Plants showed no new growth and any remaining foliage was brown and shriveled. Actual mortality could not be confirmed, as a longer monitoring period would be needed to establish true mortality (Bell et al. 2008). • Stressed: Plants were obviously unhealthy, often with yellowed, drooping leaves. • Alive: Plants were green or showed signs of healthy new growth. Data Analysis I computed the total time spent per plant, as well as the mean time per method. This was converted into mean cost per plant for each method by assuming a pay rate of $10 per person-hour based upon the pay rate for a Return of the Natives “Weed Warrior.” I also created a tally of the survivability per method in Microsoft Excel and then imported that data into SPSS where I ran a chi-squared ( 2) test on the survivorship versus the method used. I calculated the cost of Rodeo used by assuming a price of $60.00 per gallon. RESULTS Control Efficacy The most successful method I tested, which resulted in 100% apparent mortality, was digging the individual plants out. Chop and spray was a close second with 96% apparent mortality. The method of chopping repeatedly was ineffective, Crossosoma 35(2), Fall -Winter 2009 91 resulting in survivability very similar to that of the control plants (.. i2 - 67.706, p<.001) (Fig. 1). Survivorship of Fennel Plants Control Chop Repeatedly Chop and Spray Dig Treatment Method Fig. 1 - Cumulative Foeniculnm vulgare survivorship in each of the four treatment methods. *Two plants listed as dead in the chop repeatedly method could not be located during and after a re-treatment day on 21 August 2006. Cost The most expensive method was digging, with an average cost of S3. 49 per stressed or apparent dead plant (based on estimated labor cost of $10 per hour per person). The least expensive was chop and spray, with an average labor cost for two people of $0.16 per plant ($0.08 per person per plant) and an average herbicide cost of less than one cent (0.79) per plant. Chopping repeatedly was also fairly inexpensive (Table 1 ). These costs only include time spent at each plant and excludes all travel and preparation time. Table 1 - Cost average and range per-plant, by treatment method. Chop Repeatedly Chop and Spray Dig Average Cost $0.22 $0.16 $3.49 Minimum Cost $0.06 $0.06 $0.36 Maximum Cost $0.77 $0.31 $11.33 92 Crossosonui 35(2), Fall -Winter 2009 DISCUSSION Chopping and spraying each Foeniculum vulgare plant with herbicide was the least expensive and second most effective method, even with the added cost of Rodeo and the labor costs for two people. Herbicide use was determined by subtracting the amount of the solution remaining from the initial amount. Rodeo costs US $50.00 - $60.00 per gallon, depending on the source. Assuming a cost of $60.00 per gallon, the cost of the herbicide used was about 79/100 of a cent, or less than a penny, per plant. Though Roundup is a more common glyphosate formulation than Rodeo, 1 chose Rodeo because it was already available through Return of the Natives. Furthermore, Rodeo is preferable for use in riparian or aquatic sites because the surfactant in Roundup (and often added to Rodeo) has shown deleterious effects upon amphibians (Relyea 2005; Trumbo 2005). All herbicide applications for this work were on upland sites and no surfactants were used. Digging was the most effective method of controlling F. vulgare , but was labor intensive, averaging about 20 minutes for each plant. One notable plant took 68 minutes to dig out. The time may be reduced by using a different tool, such as a Pulaski or mattock instead of a shovel, however in my experience, it is veiy difficult to judge how much of the root has been removed with the Pulaski, since the tool is well suited to chopping away at the root but it is not very precise in digging. I used a shovel so that removal of the large, fleshy part of the root could be reliably determined. None of the plants had their entire root removed; once the large, bulbous portion of the root was dug up, fibrous parts of the root remained which I was unable to fully extract. Holloran et al. (2004) suggests that removing the top three to six inches of the root crown will kill the plant if the entire root cannot be removed. The monitoring portion of my study was too short to conclusively demonstrate if the small root fragments left behind after digging were enough to permit re-sprouting. Based upon my results, for small populations of F. vulgare 1 recommend digging up each plant when soils are soft, such as during the rainy season. During the summer, when the ground is too hard to dig up each plant quickly, I recommend chopping and spraying. Ideally, most removal of small and scattered populations of F. vulgare would take place in conditions where digging is the appropriate method. Unfortunately F. vulgare is often more visible during the summer, after it has sent up flower stalks, which necessitates a viable alternative to digging in hard soils. Crossosoma 35(2), Fall-Winter 2009 93 The tools used in this project, particularly the machete for chopping and shovel for digging, were chosen from personal experience, and for their appropriateness to the methods I was testing. Aside from the alternative digging tools already mentioned, loppers could also be used in place of the machete. The F. vulgare at my study sites were too widely spaced to justify the use of mowing, though brush cutters could be a possible alternative. However, Brenton and Klinger’s (2002) study on Santa Cruz Island found that cutting the plants before herbicide application did not lead to a greater reduction in F. vulgare , possibly because the fallen stems intercepted the spray. In comparison, my study was much smaller and focused upon widely spaced individual plants instead of plots with dense F. vulgare stands. I was able to clear the stalks and other cuttings from the cut section and apply herbicide directly to the exposed cut. The team of Bell et al. (2008) intensively studied herbicide control of F. vulgare and found more success with broadcast application as opposed to spot spray applications, though again their study site was much larger than mine and supported a denser F. vulgare infestation. Their results also showed that initial assessments of F. vulgare mortality can be misleading as the percent control decreased over time. Triclopyr and a mixture of glyphosate and triclopyr were more effective than glyphosate alone, especially low dosages of glyphosate. Flaming was discarded as a method since Erskine et al. (2005) indicated that burning the plants was only usefi.il in removing the previous year’s dead stalks and Klinger (2000) reported that burning is not an effective control method by itself. I observed that F. vulgare quickly regenerated from its root crown after fire and was one of the first plants to re-emerge in a field that burned in June 2006 next to the Natividad Creek Park site (Briscoe 2006). 1 did not find any references in the literature that documented the effect of fire upon the seed bank of F. vulgare . The disappearance of two plants in the chop repeatedly group partway through the study was particularly curious as they had been reliably located during three previous re-treatments. Anecdotally I observed a F. vulgare plant in a different part of the park yanked underground, possibly by a California pocket gopher, and it’s possible that the two plants in my study were preyed upon in a similar manner. Studies on gopher depredation of F. vulgare would be interesting. Further studies on how much root needs to be removed to prevent F. vulgare from re-sprouting and seed bank viability of F. vulgare after fire would be particularly useful. 94 Crossosoma 35(2), Fall -Winter 2009 ACKNOWLEDGMENTS I would like to thank Return of the Natives for providing the tools and materials used in the project, Deanne Gwinn for field assistance, Suzanne Worcester and Laura Lee Lienk for review of the project and early versions of the manuscript, as well as Scott White and Robert Klinger for very helpful reviews of this manuscript. LITERATURE CITED Bell, C. E., T. Easley, and K. R. Goodman. 2008. Effective fennel ( Foeniculwn vulgare) control with herbicides in natural habitats in California. Invasive Plant Science and Management 1 : 66-72. Brenton, R.K. and R.C. Klinger. 2002. Factors influencing the control of fennel ( Foeniculwn vulgare Miller) using Triclopyr on Santa Cruz Island, California, USA. Natural Areas Journal 22(2): 135-147. Briscoe, A. 2006 June 16. Winds fan fire in North Salinas. The Monterey County Herald Sect A 1 . California Invasive Plant Council. California Invasive Plant Inventory Database http: //www. cal-ipc. org/ ip/ inventory /weedl ist.php Accessed 2009 September 22. Erskine Ogden, J.A. and M. Regmanek. 2005. Recovery of native plant communities after the control of a dominant invasive plant species, Foeniculwn vulgare'. Implications for management. Biological Conservation 125: 427-439. Holloran, P, A. Mackenzie, S. Farrell, and D. Johnson. 2004. The Weed Worker’s Handbook: A Guide to Techniques for Removing Bay Area Invasive Plants. The Watershed Project and California Invasive Plant Council, Richmond, CA. Klinger, R.C. 2000. Foeniculum vulgare. Pages 198-202 in C.C. Bossard, J.A. Randall, and M.C. Hoshovsky (editors). Invasive Plants of California's Wildlands. University of California Press, Berkeley, California. Luken, J.O. and J.W. Thieret. 1997. Assessment and Management of Plant Invasions. Springer, New York, NY. Myers, J.H. and D.R. Bazely. 2003. Ecology and Control of Introduced Plants. Cambridge University Press, Cambridge, UK. Pimental. D., L. Lach, R. Zuniga, and D. Morrison. 2000. Environmental and economic costs of nonindigenous species in the United States. BioScience 50(1): 53-65. Pysek, P, D. M. Richardson, M. Rejmanek, G. L. Webster, M. Williamson, and .1. Kirschner. 2004. Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53: 131-143. Crossosoma 35(2), Fall -Winter 2009 95 Relyea, R.A. 2005. The lethal impact of Roundup on terrestrial and aquatic amphibians. Ecological Applications 15(4): 1118-1 124. Richardson. D. M, R Pysek, M. Rejmanek, M. G. Barbour, F. D. Pannetta, and C. J. West. 2000. Naturalization and invasion of alien plants: concepts and definitions. Diversity > and Distributions 6: 93-107. Trumbo, J. 2005. An assessment of the hazard of a mixture of the herbicide Rodeo and the non-ionic surfactant R-ll to aquatic invertebrates and larval amphibians. California Fish and Game. 91(1): 38-46. Weber, E. 2003. Invasive Plant Species of the World: A Reference Guide to Environmental Weeds. CABI Publishing, Cambridge, MA. 96 Crossosoma 35(2), Fall -Winter 2009 NOTEWORTHY COLLECTION RIVERSIDE COUNTY, CALIFORNIA Cynanchum Utah ease (Engelm.) Woodson (Asclepiadaceae) - Riverside County. Duncan S. Bell 273 (RSA, duplicates to be distributed) 19 April 2009. Palen/ McCoy Wilderness Area, west side of the Arica Mountains. Collections taken from sandy wash at 34.010° N, 114.939° W. ca. 1280 ft. elev. Large population of Cynanchum utahense observed in main drainage wash on the west side of the mountain range over a linear distance of about 300 m. AH plants in flower with approximately 30% of plants with both flowers and fruit. Growing with Ambrosia dumosa, Androstephium breviflorum , Hesperocallis undulata, Larrea trident at a, Loeseliastrum matthewsii, Malacothrix glabrata, Oenothera deltoides, Psorothamnus spinosus, Rafinesquia neomexicana, Stephanomeria exigua. Previous knowledge. Cynanchum utahense occurs from southwestern Utah, through the Mojave Desert across southern Nevada, western Arizona, to the Sonoran and Mojave deserts in California (Munz 1974, Flora of Southern California, UC Press, Berkeley; Cronquist et al. 1984, Intermountain Flora Vol. IV, New York Botanical Garden, Bronx, NY; McLaughlin 1993, Jour. Arizona- Nevada Acad. Sci. 27:169-187). Most collections of Cynanchum utahense in California have been from San Bernardino County; many of these are from the southwestern section of the Twentynine Palms Marine Corps Base. It is on CNPS List 4.2, that is, a plant of limited distribution (watch list), fairly endangered in California (i cnps.site.aplus.net/cgi-bin/inv/inventoty.cgi , visited 1 Nov 2009). The Consortium of California Herbaria (ucjeps.berkeley.edu/ consortium/ ', visited 1 Nov 2009) reports ten collections from San Diego County, mostly within the Anza-Borrego area, and only one collection from Imperial County, at Coyote Wells, south of Anza-Borrego Desert State Park. It reports two collections from Riverside County, one of which has incorrect location data, and was actually collected in San Diego County (K. Brandegee, April 1899, San Felipe). The only reported C. utahense collection made within Riverside County is Theo Glenn’s no. 91-84; May 8, 1991 (UCR) from the Sonoran Desert, Big Maria Mtns., north of Blythe. Except for Glenn’s one specimen, there is a wide gap in C. utahense s apparent distribution between western Arizona and all collections from other California counties. Significance. This collection documents a large population (more than 500 individuals) from a broad geographic area where only one other specimen is reported. This occurrence suggests that there may be other C. utahense populations in unexplored or under-collected desert washes and canyons. -Duncan S. Bell, Rancho Santa Ana Botanic Garden, I500N. College Avenue, Claremont, CA 9171 1 Become a Member of Southern California Botanists Today! □ Organization $35.00 □ Renewal Name Address 1 Address 2 City State Zip Code Phone (optional) Email address In addition, I would like to give $ to help support SCB. Make Checks payable to Southern California Botanists and mail to: Treasurer, Southern California Botanists Rancho Santa Ana Botanic Garden 1 500 North College Avenue Claremont, CA. 91711 N®w Y° k Botanical Garden Librai Southern California Botanists, Inc. - Founded 1927 - http://www.soca/bot. org 3 5185 00268 0336 Membership, Subscriptions, and Back Issues Individual and Family Memberships in SCB are $25 per calendar year domes- tic, and $35 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 $35 per calendar year, and $45 to foriegn in- stitutions. Back issues (Volume 35 - present) are available for $7 each, postpaid. Volumes 18-34 are available at $6 each. Prior to 1990, CROSSOSOMA included time-dated notices to the membership and was published six times a year. These back issues of Volumes 1 - 17 are $0.50 each, postpaid. Some back issues that are out of stock may be provided as photocopies. Available SCB Special Publications No. 1 A Flora of the Santa Rosa Plateau , by Earl W. Lathrop and Robert F. Thome, 30 pp $7.00 No. 3 Endangered Plant Communities of Southern California , Proceedings of the 15th Annual SCB Symposium, edited by Allan A. Schoenherr, 114 pp $12.00 No. 4 Flora and Ecology of the Santa Monica Mountains , 194 pp $40.00 The following article has been reprinted separately, with covers, and is available for plant collecting workshops: Herbarium Specimens as Documents: Purposes and General Collecting Tech- niques', by T. S. Ross [from Crossosoma 22(1): 3-39, 1996] $3.95 each; 10 for $22.50 Prices include California State sales tax, handling, and domestic postage. 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