VOLUME 59, NUMBER 4 OCTOBER—DECEMBER 2012 c “hie C2 A WEST AMERICAN JOURNAL OF BOTANY POINT OF VIEW CHANGES TO THE BOTANICAL CODE AND WHAT THEY MEAN FOR WESTERN NORTH AMERICAN BOTANY NA VOV GLOW "QING VIED IRULICT. cornetdlunedeint cate Men oale Seaiany ear cascaen heed ucla sven buGeners 169 CONTENTS CAREX ALBIDA (CYPERACEAE), AND ITS RELATIONSHIP TO CAREX LEMMONII Peter F. Zika and Barbara L. WilsSOM ....0.0. CARES occocee0c0ec0 Ghee C4 ocececesceccess 71 GENETIC STRUCTURE AND OUTCROSSING RATES IN VIOLA PEDUNCULATA (VIOLACEAE), A CALIFORNIA ENDEMIC VIOLET LACKING CLEISTOGAMOUS FLOWERS Theresa M. Culley and Richard L. StOKCS .eccccccccccccccccccccccccccececcceeeeeseeseees 18] RESPONSE OF GRASSLAND VEGETATION ON SANTA CRUZ ISLAND TO REMOVAL OF FERAL SHEEP Dirk H. Van Vuren and Lizabeth BOWEN cc.cccccccccccccuccecececuecececececcccecececccces 190 GENETIC STRUCTURE OF VACCINIUM PARVIFOLIUM (ERICACEAE) IN NORTHERN CALIFORNIA REVEALS POTENTIAL SYSTEMATIC DISTINCTIONS Jennifer De Woody, Valerie D. Hipkins, Julie Kierstead Nelson, and Len Lind strand VF, .occcccccccx MAI wav scco see MUNA ccc cesccsesceseoonaes 196 NOMENCLATURE OF SUBDIVISION WITHIN PHACELIA (BORAGINACEAE: HyYDROPHYLLOIDEAE) Genevieve K. Walden and Robert PAttersOn ....cccccccccccccseccccsecccuscccesseeuess At NEW SPECIES DUDLEYA CRASSIFOLIA (CRASSULACEAE), A NEW SPECIES FROM NORTHERN BAJA CALIFORNIA, MEXICO Mark W, Dodero and Michael Ge SUNDSON ics csiviscstehentdccvcesdoadostvevteatcereese 229 NOTEWORTHY (CIS TETITRCO) Pen TT aah or Na ARR cRORe eRT ee nn k SISA ee EOREE S T OP gS 230 COLLECTIONS COMETS TONG. ee ae nN eae ET aa a tT oe rene 231 (CO7AI BUTT) QIN Bs Care een 2 UT ge RM RP ESTO ACS ICES tt URN amen REPENS TEER C/N Ss de ana RT es ar SOLO ne mA SR =) ROE ee Ee ne RTO eden On 232 PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. POSTMASTER: Send address changes to MADRONO, Kim Kersh, Membership Chair, Uni- versity and Jepson Herbarium, University of California, Berkeley, CA 94720-2465. kersh @berkeley.edu. Corresponding Editor—MatT RITTER Copy Editor—RICHARD WHITKUS Biological Sciences Department Department of Biology Cal Poly, San Luis Obispo Sonoma State University 1 Grand Avenue 1801 E. Cotati Avenue San Luis Obispo, CA 93407 Rohnert Park, CA 94928-3609 madronoeditor @ gmail.com whitkus @sonoma.edu Editorial Assistant—DYLAN NEUBAUER Book Editor—MaAatTT RITTER Noteworthy Collections Editor—DIETER WILKEN Board of Editors Class of: 2012—GRETCHEN LEBUMN, San Francisco State University, CA ROBERT PATTERSON, San Francisco State University, CA 2013—ErIc ROALSON, Washington State University, WA KRISTINA SCHIERENBECK, California State University, Chico, CA 2014—BRANDON PRATT, California State University, Bakersfield, CA Tom WENDT, University of Texas, Austin, TX CALIFORNIA BOTANICAL SOCIETY, INC. OFFICERS FOR 2011-2012 President: V. Thomas Parker, Department of Biology, San Francisco State University, San Francisco, CA 94132, parker @sfsu.edu First Vice President: Andrew Doran, University and Jepson Herbaria, University of California, Berkeley, CA 94720, andrewdoran @ berkeley.edu Second Vice President: Vacant. Recording Secretary: Michael Vasey, Department of Biology, San Francisco State University, San Francisco, CA 94132, mvasey @sfsu.edu Corresponding Secretary: Anna Larsen, Jepson Herbarium, University of California, Berkeley, CA 94720, secretary @calbotsoc.org Treasurer: Thomas Schweich, California Botanical Society, Jepson Herbarium, University of California, Berkeley, CA 94720, tomas @schweich.com The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, Dean Kelch, Jepson Herbarium, University of California, Berkeley, CA 94720, dkelch@berkeley.edu; the Membership Chair, Kim Kersh, University and Jepson Herbaria, University of California, Berkeley, CA 94720, kersh @berkeley.edu; the Editor of Madrofio; and three elected Council Members: Chelsea Specht, Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720-2465, cdspecht @berkeley.edu; Ellen Simms, Department of Intergrative Biology, 1005 Valley Life Sciences Bldg., #3140, University of California, Berkeley, CA 94720, esimms @berkeley.edu. Staci Markos, University and Jepson Herbaria, University of California, Berkeley, CA 94720, smarkos @berkeley.edu. Graduate Student Representatives: Genevieve Walden, Department of Integrative Biology and Jepson Herbarium, University of California, Berkeley, CA 94720, gkwalden @ gmail.com. Administrator: Lynn Yamashita, University of California, Berkeley, CA 94720, admin @calbotsoc.org. Webmaster: Ekaphan (Bier) Kraichak, University of California, Berkeley, CA 94720, ekraichak @ gmail.com. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MADRONO, Vol. 59, No. 4, pp. 169-170, 2012 POINT-OF-VIEW — CHANGES TO THE BOTANICAL CODE AND WHAT THEY MEAN FOR WESTERN NORTH AMERICAN BOTANY The modern botanical code is the result of decisions and compromises made by numerous individuals and committees over the past several hundred years. Its purpose is to create a standard for the conventions applied to the naming of algae, fungi, and plants. The 18'" meeting of the International Botanical Congress (IBC) took place in Melbourne, Australia during the summer of 2011, where several important amendments were made to the code. One change resulted in the renaming of the code itself. To reflect a more accurate understanding of the set of rules governed by the Nomenclature Section of the IBC and its voting body, the /nternational Code of Botanical Nomenclature (ICBN) will now be known as the International Code of Nomenclature for algae, fungi, and plants (ICN) (McNeill and Turland 2011). The Nomenclature Section of the IBC in Melbourne also ruled that new species diagnosis and/or descriptions may now be written in English. Previously, to validly publish a new name, a diagnosis (how the new plant differs from its close relatives) and/or description (a comprehensive summary of the characters) in Latin was required to introduce the taxa into science. Article 36 in the revised code states that as of January 1, 2012, publication of a new species does not have to be written in Latin, but can be replaced with English (Smith and Figueir- edo 2011). Ultimately, the decision of whether to write the description in English or Latin rests with the journal or publisher (Knapp et al. 2011). This change will not affect the scientific naming of plants, which will still be done in Latin. Articles 29, 30, and 31 of the updated code outline the use of online publishing as an accepted way to publish new plant names and name changes. Prior to this change, a printed version was required to validly publish a new species name. According to Article 29, an online publication accompanied by either an Interna- tional Standard Serial Number (ISSN) or an International Standard Book Number (ISBN) in a PDF format will suffice (Knapp et al. 2011). The articles also list several recommendations such as depositing printed material of new names or name changes in libraries on several continents and publishing in journals that are archived in several online repositories. Article 30 states that electronic publishing will not be recognized if the publication date was before January 1, 2012, and indication of draft and final versions of publications should be clear. Article 31 states that eit publications of printed and online material must be treated as having the same date, and that the publication date should be clearly stated somewhere in the article. If the PDF format becomes outdated and another format becomes more widespread, the ICB allows the terms of best practice to conform to these changes. These updated methods and recommen- dations will help botanists publish and distribute new botanical information in a timely, standard- ized, and efficient manner. The most publicized and controversial decision made by the Nomenclature section of the IBC was over the retypification of the genus Acacia, which has been shown to be polyphyletic and therefore in need of revision (Luckow et al. 2003). According to the code, if a genus is split, the type specimen may be changed if the split causes too much disruption and renaming of species. During the 2005 Vienna conference, the ruling was made to change the type specimen from A. scorpioides (L.) W. Wight (formerly A. nilotica Karst.) to A. penninervis Sieber ex DC. The retypification conserved 960 species of Acacia, and caused 160-170 to be placed in the separate genera including: Acaciella, Senegalia, and Vachellia (Luckow et al. 2005). For California botanists, our native catclaw (Acacia greggii) was renamed Senegalia greggii (Baldwin et al. 2012). The conservation of the genus allowed most of Australia’s wattles to remain in Acacia, while acacias in Asia, the Americas, and especially Africa required name changes. Several botanists challenged the ruling, believing it to be unfair particularly to African botanists (Moore and Cotterill 2011). These changes and revisions make publishing new species more efficient and accessible to the next generation of botanists. Publishing new names and name changes online, along with allowing new species descriptions in English, will help modernize the way botanists work. Although the changes made to Acacia were controversial, the conservation of the name was with the most speciose group in the former genus, and the change helps us better understand the complexity of this large group of trees and shrubs. —Taylor Crow and Matt Ritter, Biology Department, Cal Poly, San Luis Obispo, CA 93407, tmcrow8@gmail.com LITERATURE CITED BALDWIN, B. G., D. H. GOLDMAN, D. J. KEIL, R. PATTERSON, T. J. ROSATTI, AND D. H. WILKEN, (eds.). 2012. The Jepson manual: vascular plants of 170 MADRONO [Vol. 59 California, second edition. University of California Press, Berkeley, CA. KNappP, S., J. MCNEILL, AND N. J. TURLAND. 2011. Changes to publication requirements made at the XVIII international botanical congress in Melbourne—what does E-publication mean for you? Taxon 60:1498—1501. Luckow, M., J. T. MILLER, D. J. MURPHY, AND T. LIVSHULTZ. 2003. A phylogenetic analysis of the Mimosideae (Leguminosae) based on chloroplast DNA sequence data. Pp. 197-220 in B. B. Klit- gaard and A. Bruneau, (eds.), Advances in Legume systematics, part 10. Higher level systematics. Royal Botanical Gardens, Kew, U.K. , C. HUGHES, B. SCHRIRE, P. WINTER, C. FAGG, R. FORTUNATO, J. HURTER, L. Rico, F. J. BRETELER, A. BRUNEAU, M. CACCAVARI, L. CRAVEN, M. Crisp, A. DELGADO S, S. DEMISSEW, J. J. DOYLE, R. GRETHER, S. HARRIS, P. S. HERENDEEN, H. M. HERNANDEZ, A. M. HIRSCH, R. JOBSON, B. B. KLITGAARD, J.-N. LABAT, M. Lock, B. MACKINDER, B. PFEIL, B. B. StIMPSON, G. F. SMITH, M. SOUSA S., J. TIMBERLAKE, J. G. VAN DER MAESEN, A. E. VAN Wyk, P. VORSTER, C. K. WILLIS, J. J. WIER- INGA, AND M. F. WOJCIECHOWSKI. 2005. Acacia: the case against moving the type to Australia. Taxon 54:513—519. MCNEILL, J. AND N. J. TURLAND. 2011. Major changes to the Code of Nomenclature—Mel- bourne, July 2011. Taxon 60:1495—-1497. Moore, A. AND F. COTTERILL. 2011. The Acacia retypification debate: perspectives of African am- ateur botanists. Taxon 60:858—859. SMITH, G. F., E. FIGUEIREDO, AND G. MOORE. 2011. English and Latin as alternative languages for validating the names of organisms covered by the International Code of Nomenclature for algae, fungi, and plants: the final chapter? Taxon 60:1502—1503. MADRONO, Vol. 59, No. 4, pp. 171-180, 2012 CAREX ALBIDA (CYPERACEAE), AND ITS RELATIONSHIP TO CAREX LEMMONII PETER F. ZIKA WTU Herbarium, Box 355325, University of Washington, Seattle, WA 98195-5325 Zikap@comcast.net BARBARA L. WILSON Carex Working Group, 1377 SW 13" Street, Corvallis, OR 97330 ABSTRACT Carex lemmonii W. Boott is a widespread California endemic. Herbarium and field studies showed its extensive morphological variation included plants called Carex albida L. H. Bailey, previously considered endemic to Sonoma County, California. Due to extensive overlap, characters of the perigynium, achene, inflorescence, and foliage did not separate the two taxa. Carex albida becomes a synonym of the earlier name, Carex lemmonii, which has implications beyond nomenclature. Carex lemmonii is common and is not a conservation priority in California. When merged with Carex lemmonii, the only known natural population of Carex albida will no longer qualify for protection under federal and state endangered species legislation. A key and illustrations are provided for California species in Carex section Au/ocystis Dumort. Key Words: California, Carex albida, Carex lemmonii, Cyperaceae, endangered species. Carex sect. Aulocystis Dumort. (syn. sect. Ferrugineae (Tuckerman) Kuk.) is represented by eight species in North America, with five species recorded in California (Ball and Mastro- giuseppe 2002). One of these is C. albida L. H. Bailey, white sedge, restricted to Sonoma County, California. It was first collected on 1 May 1854, by Dr. Jacob M. Bigelow, expedition surgeon and botanist for Lieutenant Amiel W. Whipple’s exploration for a railway route from the Muis- sissipp1 River to the Pacific Ocean, near the 35th parallel of latitude (Torrey and Gray 1857, p. 154). References to George Thurber as a collector of C. albida (e.g., Boott 1880; Bailey 1889; Kukenthal 1909) are apparently an error, according to Howell (1957). Thus Bigelow was the only botanist to find C. albida in the 1800's, perhaps from present day Laguna de Santa Rosa (Best et al. 1996). There are only a couple additional locations from the 1900’s (Howell 1957). At present, plants called Carex albida are reduced to a single extant population in the diminished remnants of Pitkin Marsh wetlands, at an elevation of 30 m. The same site also supports a population of the Federally Endan- gered Lilium pardalinum Kellogg subsp. pitki- nense (Beane & Vollmer) Skinner. Carex albida received protection as State Endangered under California legislation in November 1979. Federal listing as an Endangered Species came 22 October 1997 (62 FR 55791). Carex lemmonii W. Boott, Lemmon’s sedge, as traditionally defined, is a California endemic, known from 25 counties across the state, ranging in elevation from 700 to 3000 m (Consortium of California Herbaria 2010). It favors creek banks, wet meadows, boggy meadows, fens, marshes, and peatlands, and occasionally is found on serpentine soils. The type material of Carex albida was 1mma- ture, and for many decades there were no additional collections. Eventually botanists ques- tioned its taxonomic validity. Mackenzie (1922) combined C. alhida with C. luzulina Olney. Later, Mackenzie (1935, 1940) lumped C. albida with C. lemmonii. However, his broad circumscription of C. lemmonii was not followed by Stacey (1937), who described C. albida again, as C. sonomensis Stacey. Howell (1957, 1965) included C. sono- mensis within C. albida, while retaining C. lemmonii as a distinct species. His approach was followed by Mastrogiuseppe (1993) and then Ball and Mastrogiuseppe (2002). These different taxonomies were difficult to reconcile when preparing a treatment of Carex for the revision of the Jepson Manual (Mastrogiuseppe 1993; Zika et al. 2012). The purpose of this study was to evaluate the morphological characters various authors have used to discriminate between C. albida and C. lemmonii, and explain the revised classification presented here. METHODS The sedge literature was reviewed for morpho- logical characters that separate Carex albida from C. lemmonii. The taxonomic concepts of the most recent treatment, Ball and Mastrogiuseppe (2002) served as a starting point. Characters of foliage, perigynia, achenes and inflorescences used in the literature over the last century were measured on eZ TABLE 1. MADRONO [Vol. 59 A COMPARISON OF CAREY ALBIDA (INCLUDING THE TYPE OF C. SONOMENSIS) AND C. LEMMONII (INCLUDING THE TYPE OF C. ABRAMSII), WITH EMPHASIS ON CHARACTERS HISTORICALLY USED TO DISTINGUISH BETWEEN THE Two TAXA. All specimens examined for C. a/bida are from Sonoma Co. (n = 39). All entries for the C. lemmonii column are from outside Sonoma County (n = 270). Achene length includes the stubby stipe at the base of the achene body, but does not include the irregular style remnants above the achene body. Taxon Character C. albida (Sonoma Co.) Dried leaf width (mm) be -35.6 Fresh leaf width (mm) 2-8 (cultivated) Inflorescence length (cm) 2.5—37 Inflorescence lowest 1=19 internode (cm) Distal lateral spikes Yor N clustered Terminal spike overtops Y, or slightly lateral spikes Sex of terminal spike (S = staminate; P = pistillate) Lateral spike length (mm) Scale color outside the green midrib S, S/P, S/P/S, P/S, P 5-15 white, green, light brown, gold, sometimes red- brown splotches Perigynium color green, light brown Perigynia (mm) 3.1-4.2 * 1.0-1.5(1.8) Perigynium beak teeth Yor N bristly Achene outline obovoid, obovoid-elliptic, obovoid-oblong, ovate- elliptic, ovate-oblong, elliptic Achene (mm) 1.6-1.9 X 1.05—1.2 herbarium specimens, using a micrometer and dissecting binocular microscope. Gatherings of C. albida (as well as its synonym C. sonomensis) and C. /emmonii (as well as its synonym C. abramsii Mack.), were studied at the following institutions: BM, CAS, CHSC, DS, F, G, GH, HSC, JEPS, K, MICH, MO, NY, ORE, OSC, NY, POM, RSA, S, UC, US, WILLU, WTU, and YM (acronyms from Holmgren et al. 1990). All available types were scrutinized. This preliminary study utilized 39 herbarium speci- mens of C. al/bida, and 270 sheets of C. lemmoniti. The range of variation for 13 characters, across all of California, was summarized in Table 1. A principal components analysis (PCA) was performed in the R environment for statistical computing (R Development Core Team 2010). Ordination was performed using Euclidian dis- tances. Non-metric multidimensional scaling C. lemmonii (except Sonoma Co.) 1.2-5.8 1.5—6.5 (wild) 5—49.2 1 .5—20 YorN Y, or slightly S, S/P, S/P/S, P/S, P 6—22 white, green, light brown, gold, red-brown, sometimes dark purple splotches green, light brown, sometimes red-brown or dark purple splotches (2.5)2.8-4.5(4.8) x (0.8)1.0—1.5(1.8) YorN Author using character Mastrogiuseppe (1993); Ball and Mastrogiuseppe (2002) Mastrogiuseppe (1993); Ball and Mastrogiuseppe (2002) Mastrogiuseppe (1993) Kiukenthal (1909); Stacey (1937); Howell (1957, 1965); Mason (1969) Stacey (1937) Stacey (1937); Howell (1965); Mason (1969); Mastrogiuseppe (1993) Mastrogiuseppe (1993) Kiukenthal (1909); Stacey (1937); Howell (1957, 1965); Mason (1969) Howell (1957) Howell (1965); Mastrogiuseppe (1993) Howell (1957) obovoid, obovoid-elliptic, Stacey (1937) obovoid-oblong, ovate- elliptic, elliptic, broadly elliptic 1.42.0 < 0.8-1.3 US Fish and Wildlife Service (2009) (NMS) was performed using PC-ORD (McCune and Mefford 2006), using Sorenson distances. The maximum number of iterations for the algorithm to perform was set to 200, however, the analysis reach a low-stress (10.25%) plateau after 83 iterations were performed. ANOVA, NMS, and PCA were based on a subset of 64 herbarium specimens, evaluating 18 morpholog- ical characters (Table 2). The vouchers are cited in Appendix 1; they included six isotypes, seven specimens of Carex albida and 57 specimens of C. lemmonii (Table 3). Sampled specimens came from 12 counties and six ecoregions in California (Baldwin et al. 2012). Field visits were made to Carex /emmonii populations in Butte, Mariposa, and San Ber- nardino counties in 2009 and 2010. Leaf width was measured fresh on ten cultivated plants of C. albida at the Botanical Gardens of the University 2012] TABLE 2. THE 18 MORPHOLOGICAL CHARACTERS EVALUATED FOR ANOVA, PCA AND NMS ANALYSES. Morphological character Width of narrowest basal leaf (mm) Width of widest basal leaf (mm) Stem height (cm) Inflorescence length (cm) Length of proximal inflorescence internode (cm) Clustering of distal spikes (yes or no) Distance from apex of distal lateral spike to apex of terminal spike (mm) Sex of terminal spike Length of shortest lateral spike (mm) Length of longest lateral spike (mm) Color of pistillate scales (whitish-green, red-brown, dark purple) Color of perigynia (whitish-green, red-brown, dark purple) Length of perigynium (mm) Width of perigynium (mm) Perigynium beak teeth with bristles (yes or no) Distance from achene apex to perigynium beak apex (mm) Length of achene body (not including stipe or beak; mm) Width of achene body (mm) of California in Berkeley, in November 2009. Corresponding measurements were made on living plants of C. /emmonii in meadows along the South Fork of the Santa Ana River in San Bernardino County in August 2010. Vouchers of C. lemmonii were deposited at CHSC, JEPS, MICH, OSC, RSA, and WTU. RESULTS Taxonomists often use perigynium differences to separate closely related Carex species. Photo- graphs of the perigynia of each California member of sect. Au/ocystis are shown in Fig. 1. Typical C. lemmonii perigynia (Fig. 1|A—E) show TABLE 3. ZIKA AND WILSON: CAREX LEMMONII 173 variation in length, width, outline, color, streaking, and marginal bristles. Each sample of 4—5 perigynia was derived from a single herbarium specimen. With practice, C. /emmonii perigynia are distin- guishable from C. /uzulina (Fig. 1G) by beak length and color differences. Carex luzulifolia S. Watson perigynia (Fig. 1F) have broader wings and longer, narrower, more abrupt beaks than C. lemmonii. Carex fissuricola Mack., with perigynia somewhat intermediate between C. /uzulina and C. luzulifolia, is separable by its stiff spreading- ascending bristles just visible as a sparse white fuzz (Fig. 1H). In summary, typical perigynia of C. luzulina, C. luzulifolia, and C. fissuricola look different from typical perigynia of C. lemmonii, as shown in Fig. |. But within C. /emmonii plants from Sonoma Co. (Fig. 1A) are not distin- guishable from specimens from Mariposa Co. (Fig. 1B). Perigynia from the types of C. sono- mensis (Fig. 1A) and C. abramsii (Fig. 1E) merge imperceptibly with perigynia of C. /emmonii from elsewhere across the California floristic province. This pattern of overlap was also true for the other features measured. A detailed examination of critical morphological characters, including foliage, inflorescences, spikes, scales, achenes, and perigynia, is collated in Table |. The preliminary study did not find any character that discrimi- nates between Carex albida and C. lemmonii. The most extreme difference was leaf width, which was reported to be slightly greater in C. a/bida than in C. lemmonii (Mastrogiuseppe 1993; Ball and Mastrogiuseppe 2002). Leaf width was found to decrease when specimens were dried, and was too variable between individuals to be useful (Table 1). Even with fertilizer treatments, several cultivated plants of C. albida had widest (fresh) leaves 5—5.5 mm, while other garden plants had fresh leaves 7-8 mm wide. Within this continuum fit wild plants in the San Bernardino Mountains, with widest fresh leaves 4—6.5 mm wide. Not all GEOGRAPHIC ORIGIN AND NUMBER OF HERBARIUM SPECIMENS OF CAREX LEMMONII AND ITS SYNONYMS FROM CALIFORNIA EXAMINED FOR ANOVA, PCA AND NMS ANALYSES IN THIS STUDY. See Appendix for a list of vouchers. County distribution: AL = Alpine Co., BU = Butte Co., EL = El Dorado Co., FR = Fresno Co., MA = Mariposa Co., NE = Nevada Co., PL = Plumas Co., SA = San Bernardino Co., SI = Sierra Co., SO = Sonoma Co., TU = Tulare Co., YU = Yuba Co. Ecoregion distribution (Baldwin et al. 2012): CaR = Cascade Range, NCoRO = Outer North Coast Ranges, n SN = northern Sierra Nevada, c SN = central Sierra Nevada, s SN = southern Sierra Nevada, SnBr = San Bernardino Mountains. Carex AL BU EL C. abramsii isotype C. sonomensis isotypes C. lemmonii l 23 1 CaR NCoRO nSN All Carex combined 13 7 3] California counties FR MA NE PL SA SI SO TU YU l pi 3 2 16 4 l 2 | 2 California ecoregions cSN sSN_ SnBr 5 cS) 5 174 MADRONO Fic. 1. Representative perigynia for Carex sect. Aulocystis, all specimens from California. A. Carex lemmonii from Sonoma County, an isotype of C. sonomensis (J. T. Howell & Stacey 13042). B. Carex lemmonii from Mariposa County (Bolander 4995). As in A, note large pale perigynia. C-D. Carex lemmonii from two different populations in Butte County. C. (Copeland s.n.). D. (Janeway 3039 & Schlising). E. Carex lemmonii from San Bernardino County, an isotype of C. abramsii (Abrams 2816). Note some dark purple blotches. F. Carex luzulifolia from Tuolumne County (Col/vell 5-361& Grossenbacher). Note broad flat wing around achene and narrow straight- sided beak. G. Carex luzulina from Del Norte County (Tracy 19177). Note beaks slightly longer and color slightly darker than C. /emmonii. H. Carex fissuricola from Fresno County (J. T. Howell 22451). Note pale bristles on distal perigynium surfaces. plants from Sonoma County have broad leaf blades. Given the variation within populations, we did not consider leaf width to have any practical taxonomic significance between popula- tions. Additional statistical analysis was used to address this question, as discussed below. Field visits to Carex lemmonii populations from Butte, Mariposa, and San Bernardino counties showed broad variation in many char- acters. A number of individuals were observed in the field that closely resembled herbarium spec- imens as well as cultivated plants of C. albida, in habit, and in technical details (Table 1). Quantitative Analysis An ANOVA of quantitative morphological traits found that Carex albida specimens did not 2012] ZIKA AND WILSON: CAREX LEMMONII iS TABLE 4. SELECTED QUANTITATIVE MORPHOLOGICAL TRAITS IN CAREX ALBIDA (N = 7) AND C. LEMMONII (N = 57). All C. albida specimens are from Sonoma Co., all C. /emmonii specimens are from other counties in California. C. albida C. lemmonii Trait (unit of measurement) mean (s.e.) range mean (s.e.) range Leaf width, minimum (mm) 2.6 (0.16) 2.0—3.1 1.8 (0.06) 0.9-2.8 Leaf width, maximum (mm) 4.8 (0.42) 3.2-5.9 3.2 (0.10) |e sae Culm height (cm) 62.8 (4.15) 44.9-73.3 60.5 (2.64) 27.7-114.7 Inflorescence length (cm) 19.3 (2.57) 12.0—30.0 17.5 (1.20) 5.6—52.8 Lowest inf] internode length (cm) 14.0 (2.00) 9,224.8 10.5 (0.65) 225-24 Distance between distal-most 2 spikes (mm) 10.1 (1.50) 6.0—8.0 8.3 (0.39) 2.0—17.0 Lateral spike length, minimum (mm) 7.3 (0.47) 6.09.0 7.7 (0.25) 4.0—12.0 Lateral spike length, maximum (mm) 11.3 (1.30) 8.0—17.0 12.0 (0.41) 7.0—20.0 Perigynium length (mm) 3.7 (0.08) 3.44.0 3.5 (0.05) 2.8—4.5 Perigynium width (mm) 1.3 (0.03) 1.2-1.4 1.3 (0.03) 0.9-1.7 Perigynium beak length (mm) 1.6 (0.08) 1.5—1.7 1.5 (0.04) 0.92.2 Achene length (mm) 1.5 (0.06) 1.4-1.8 1.5 (0.02) 1.2-1.8 Achene width (mm) 1.1 (0.02) 1.1-1.2 1.1 (0.02) 0.7—1.4 Perigynium length/width 2.8 (0.10) 2.5—3.2 2.8 (0.06) 1.9-3.8 Achene length/width 1.3 (0.05) 1.2—-1.6 1.4 (0.03) 1.11.8 Beak lenth/perigynium length 0.4 (0.012) 0.40.5 0.4 (0.01) 0.3—-0.5 differ from C. /emmonii specimens (F = 0.2732, P = 0.9999). Except for leaf width, measured morphological traits of the seven Sonoma spec- imens were within the range for the 57 measured C. lemmonii specimens (Table 4). The widest leaves observed were found in C. albida speci- mens and if that trait were considered alone the difference between the taxa would be considered significant (t = 3.721 and 4.979 for minimum and maximum leaf width respectively, P = 0.001 and 0.007, respectively). But there was considerable overlap of leaf width with C. /emmonii, as discussed above (Tables 1, 4). Green scale color was more common in C. albida than in C. lemmonii, but occurred in both (Tables 1, 5). Gynecandrous terminal spikes were observed in three of the seven C. alhida and not in the C. lemmonii specimens, but the other arrangements observed in C. albida were found in C. lemmonii (Table 5). Our more extensive sample of C. lemmonii did yield occasional gynecandrous terminal spikes (Table 1), so we do not believe TABLE 5. SELECTED QUALITATIVE TRAITS IN CAREX ALBIDA (N = 7) AND C. LEMMONII (N = 57). All C. albida specimens are from Sonoma Co., all C. /emmonii specimens are from other counties in California (S = staminate portion of spike; P = pistillate portion of spike). C. albida C. lemmonii C. albida C. lemmonii Trait number number percent percent Scale color green 6 23 86% 40% red-brown ] 24 14% 42% purple 0 10 0% 18% Terminal spikes clustered 6 42 86% 74% not clustered i 15 14% 26% Terminal spikes staminate Z 48 29% 84% S/P l 8 14% 14% S/P/S ] | 14% 2% P/S 3 0 43% 0% Perigynium color green 6 29 86% 51% red-brown 1 8 14% 14% purple 0 20 0% 35% Perigynium beak teeth bristly no 3 30 43% 53% yes 4 27 57% 47% 176 MADRONO N a 36 < LM and SVDE, and at locus VCC1_J9 in populations 2HC, 2SR, 3MP, and 4CC (Supplemental Mate- rial). The frequency of the detected null ranged from low (0.071) to moderate (0.262), indicating that null alleles may have inflated fixation indices (indicating a deficit of heterozygotes) in these collections. Genotypes adjusted for null alleles at each affected locus were randomized within each population and used for population-level analy- ses of differentiation and genetic distance. Analysis of molecular variance over the null- corrected data revealed significant differentiation among populations and categories of populations in this collection of Vaccinium. The one-level 2012] model estimated 58% of the variance to be contained within populations and 42% of the genetic variance to be partitioned among popu- lations (P < 0.001). Significant differentiation was observed in the two-level hierarchical mo- del based on geographic location: 51% of the variance was contained within populations, 10% of the variance was distributed among popula- tions within a region, and 39% among geographic regions (P < 0.001). Estimates of differentiation from the raw genotypic data were congruent to those from null-corrected data (data not report- ed), indicating the occurrence of null alleles in the data set was insufficient to bias the detection of differentiation among these populations. Levels of genetic distance calculated from the corrected genotypes varied among pairs of populations (Table 3). Genetic distances were greater between populations in different geo- graphical regions, and smaller among popula- tions within geographic areas. This pattern was confirmed by tests for isolation by distance (IBD), which identified a significant positive correlation between genetic and geographic distance among all pairs of populations for both raw (Rxy = 0.38, P < 0.001) and null-corrected data (Rxy = 0.39, P = 0.003). This pattern is consistent with the AMOVA analyses indicating greater differences among regions than among populations within a region. Population phenograms built from the raw or null-corrected data were mostly congruent in their topology but neither was. statistically significant (all branches were observed in less than 60% of bootstrap replicates). In both phenograms, populations grouped by geography, with the coastal populations (categories 2 and 3) forming one clade, the Sierra Nevada populations (category 4) a second clade, and the Shasta County populations (category 1) a third clade (Fig. 3). Potentially the most informative differ- ence between the raw and null-corrected pheno- grams involved the placement of the single population of V. deliciosum (SVDE). In the raw data phenogram, 5VDE is placed between the coastal clade and the Shasta and Sierra clades, while in the null-corrected data, 5VDE is placed between the Shasta and Sierra clades (Fig. 3). Together, these phenograms indicate the atypical populations from Shasta County and the central Sierra Nevada are more similar to the congener V. deliciosum than the coastal populations of V. parvifolium. While the phenograms represent the similari- ties among population averages, the Principle Coordinate Analysis (PCoA) maximizes differ- ences among individuals. The PCoA based on the raw genotypes clustered samples by geogra- phic collection and morphology on the first two axes, which explained 61% of the variation among individuals (Fig. 4). Four putative genetic DEWoopy ET AL.: GENETIC STRUCTURE OF VACCINIUM PARVIFOLIUM 203 clusters were indicated by the PCoA: V. parvifo- lium of typical morphology from the coastal areas, Vaccinium from the Sierra Nevada Moun- tains, Vaccinium from Shasta County, and the congener V. deliciosum. The first axis distinguish- es the typical V. parvifolium from the Coastal Ranges and western Oregon from the collections of atypical Vaccinium from Shasta County and the Sierra Nevada. The single population of V. deliciosum 1s intermediate to these, but more similar to the atypical Vaccinium collections (Fig. 4). The second axis further separates the atypical Vaccinium collections by geographic region. The admixture analyses using Structure esti- mated the most likely number of genetic clusters in the Vaccinium data set to be five (dK5 = 1, all other dK =~ QO). Individual assignment to each cluster roughly followed population categories per Table 1. The populations of V. parvifolium of typical morphology were genetically similar and potentially admixed, being assigned to two clusters in varying proportions (orange and red). The five populations of atypical morphology from Shasta County were assigned to a single cluster (blue). The atypical populations Vaccin- ium from the Sierra Nevada were assigned to another cluster (purple), with some admixed with the blue cluster. The single population of V. deliciosum (SVDE) was distinct and assigned to a unique genetic cluster (green) (Fig. 5). These individual assignments are concordant with the PCOoA results. The Neighbor-Joining phenogram of anony- mous genetic clusters identified in the Structure analysis was concordant with the PCoA and population phenograms. The cluster containing atypical Shasta County samples (blue) was more similar to the cluster containing atypical central Sierra Nevada collections (purple) and V. deli- ciosum (green) than to the clusters containing samples of typical morphologies (orange and red) (Fig. 5). These results are likely not an artifact of the null alleles as nulls were accounted for in the Structure analyses. Together, these results indi- cated the genetic differences among populations corresponded to geographic and morphological factors. DISCUSSION Genetic Differentiation among Sampled Populations The primary goal of this study was to establish whether populations of Vaccinium from Shasta County and the Sierra Nevada displaying atyp- ical berry color and morphology were geneti- cally similar to typical V. parvifolium from northwestern California and western Oregon and Washington. 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ACAS OWr HOr OOr Odr dINE I€ USZ WHASZ U1z az STZ GHz SHE HOVE a1n! OOSI a1 WIA Hal HdAS OWP HOP 204 ‘sorouanbaly dJa][V Po1dd1109-]]NU WIOI] [PUOSRIP 9AOQe SoN[eA Seep MPI WOLJ 1B [BUOSPIP MOTAq SON[VA “[ I[QUL, MOT[OJ SUONIADIQGY ‘ddS WAINIDIVA AO SNOILLV1NdOd OT ONOWY AONVILSIC SILANAD (ZTL61) SAAN JO SALVINILSY °¢ ATAVL$ 2012] 2HC 2SFM 2JS FIG. 3. DEWoopy ET AL.: GENETIC STRUCTURE OF VACCINIUM PARVIFOLIUM 2ACF 5Hp 2LD 2LR 2ACF 3MP 2SR Unrooted Neighbor-Joining consensus population phenograms from Nei’s (1972) genetic distances among 20 populations of Vaccinium. (A) The phenogram built from the raw data. (B) The phenogram built from null- corrected allele frequency data. Low statistical support was found for the topologies, with all branches occurring in <60% of bootstrap replicates. loci revealed significant structure and differentia- tion along geographic and morphological groups. Four statistical analyses provide congruent evi- dence that the atypical populations of Vaccinium are genetically distinct from the V. parvifolium located in the Pacific coastal areas, and are more similar genetically to V. deliciosum, indicating additional systematic consideration may be war- ranted. The level of differentiation among regions in this study was greater than the among-population variation reported for other Vaccinium species. The level of differentiation observed among populations within geographic regions in this study (10%) is consistent with the other Vaccin- ium reports (Yakimowski and Eckert 2008; Bell et al. 2009; Debnath 2009). Levels of differenti- ation among geographic regions was much higher (39%), indicating gene flow is restricted between the sampling categories (Table 1). Varying levels of vegetative reproduction were identified across the collection. Vaccinium species 206 Typical V. parvifolium A Axis 2 (17%) MADRONO V. deliciosum [Vol. 59 Atypical Shasta County Pop. la @1BH © 1FLM A1LB @1SQC x 1ULB m@ 2ACF X 2HC @2HD A2JS @2LR X2SFM = 2SR + 3MP @4BC @4CC 4 4GH =4MC A5VDE B Atypical Sierra AA - b Nevada Axis 1 (44%) Fic. 4. Principal coordinate analysis reveals genetic structure among populations of Vaccinium. Each point represents one genet, and symbols correspond to population. Color relates to morphological variation depicted in Fig. 1. are widely reported to reproduce clonally (Kreher et al. 2000; Yakimowski and Eckert 2008; Bell et al. 2009). Vaccinium parvifolium is reported to typically reproduce sexually but capable of producing sprouts in response to disturbance or damage (Wender et al. 2004) and in some cases via rhizomes (Gehrung 2001). Here, most popu- lations displayed high levels of genet diversity (>0.8), but the collections from the Sierra Nevada displayed greater vegetative reproduction than those from the Coastal areas as indicated by smaller G/N ratios. Rhizomatous species may share resources among stems via a complex root system, providing a mechanism of survival in suboptimal or heterogeneous habitats (Wender et al. 2004). Thus, differences in vegetative reproduction in Vaccinium in different geograph- ic regions may reflect an adaptive response to habitat quality or disturbance. The presence of null alleles at four of the five loci may inflate measures of fixation and bias estimates of population differentiation, yet the low frequency of most nulls and their uneven distribution among populations minimized the impact on data interpretation. Null alleles are expected to be higher in frequency when primers designed for one species are used to amplify microsatellite loci in a related species, as was done in this study. However, if the lack of amplification resulted from fixed differences between V. parvifolium and the species for which the primers were designed (V. corymbosum, highbush blueberry, Boches et al. 2005), we would expect null alleles to occur at high frequency in the majority of populations. Three lines of evidence indicate the impact of the null alleles was minimal and the genetic patterns observed were robust. First, as the AMOVA conducted for both raw data and that adjusted for null alleles produced congruent values, the results are likely conservative estimates of differ- entiation among populations and regions. Sec- ond, the Neighbor-Joining trees built with both raw data and null-corrected allele frequency data showed highly concordant topologies. Third, the Structure admixture analysis and individual assignment tests were parameterized to account for low frequencies of null alleles (Falush et al. 2007) and identified genetic structure concordant to those resolved with other analyses. A high number of unique alleles were detected in various Vaccinium populations. A total of 27 alleles (26%) were only observed in a single population, with 12 of the 20 populations displaying at least one unique allele (Supplemen- tal Material). Private alleles were more frequent in the typical coastal and atypical Sierra Nevada populations than the atypical populations of Shasta County, indicating gene flow is restricted among regions. A relatively small number of migrants per generation (V.m = 4) 1s sufficient to maintain genetic similarity between demes (Hartl and Clark 2007). Restricted gene flow between the Sierra Nevada and Coast Ranges seems 2012] DEWoopy ET AL.: GENETIC STRUCTURE OF VACCINIUM PARVIFOLIUM 207 > Mean LnP(D) —@ dk Mean LnP(D) OO Ww fb) (ok > _ (e) Cc ® od) ee o) Cc Ro) y a ie) ok ie) — QO. 1BH 1FLM 1LB 1SQCIULB2ACF 2HC 2HD 2JS 2LD 2LR2SFM2SR 31 3MP 4BC 4CC 4GH 4MC5VDE Population Fic. 5. Patterns of genetic differentiation revealed by admixture analyses. A) The average likelihood of analyses identified five genetic clusters (dK) in the collection. B) Assignment of individuals to the five genetic clusters. Each vertical bar represents one sampled population. Color corresponds to each genetic cluster; bars composed of multiple colors represent potentially admixed populations. Refer to Table 1 for sample sizes, which vary among populations. C) Neighbor-Joining phenogram of the genetic distances between the five genetic clusters. Colors match genetic clusters from (B). 208 reasonable given the geographic distance between regions. A lack of gene flow between the Coast Ranges and Shasta County may be more surprising. Some populations of differing mor- phology are closer geographically (e.g., 1SQC and 2SFM separated by 77 km) than are two populations of the typical phenotype (e.g., 1SQC and 3MP separated by 425 km). Yet the distant populations of similar morphology are signifi- cantly more similar genetically. The lack of private alleles in the Shasta County populations may be due to a number of demographic factors requiring additional study (e.g., inbreeding). The locus CA23F displayed fixed differences between geographic regions, with samples from the coastal areas being homozygous for a smaller allele (164 bp) and all other samples from the Sierra Nevada being homozygous for a larger allele (167 bp). Given the expected high rate of mutation at SSR loci, the lack of variation at this locus may indicate it is linked to a functional variant under strong selection. Additional study will be required to determine if fixed allelic differences correspond to any adaptive variation. Systematic Interpretations of Differentiation The second objective of this study was to determine whether any genetic differences be- tween populations might be indicative of greater systematic divergence than currently described in the taxonomic treatments. Morphologically, V. parvifolium forms a distinct clade separate from the V. deliciosum/ovalifolium complex (Vander Kloet and Dickinson 1999). Vaccinium parvifo- lium is typically larger than related Vaccinium species, and is characterized by slower germina- tion and apical meristem development in seed- lings, and persistent juvenile leaf morphology, distinctive biochemical properties and a taproot system unique in the section. Phenotypic variation within V. parvifolium is well noted (Vander Kloet and Dickinson 1999; Gehrung 2001). Gehrung (2001) noted the rhizomatous growth and distinct berry color in the Sierra Nevada populations, and suggested elevation to subspecific status may be appropriate upon further study. However, if variation is due to phenotypic plasticity among panmictic, we would expect low genetic differen- tiation between morphologically distinct popula- tions, which is not the pattern observed here. Further, phenotypic variation in the related V. membranaceum Douglas ex Torr.was shown to be environmentally induced (Schultz 1944 in Vander Kloet and Dickinson 1999); common garden experiments are required to determine the genetic basis of morphological variation in V. parvifolium. Vaccinium deliciosum, 1n contrast, is rhizomatous, forming dense stands of ramets, and displays very little phenotypic variation (Gerhung 2001). Ge- netic examinations have also separated these MADRONO [Vol. 59 species into different clades of section Myrtillus. Cytological evidence reported V. parvifolium from the Pacific Coast (British Columbia, Canada and Washington state) as diploid (7 = 12) while V. deliciosum was described as tetraploid (n = 24) (Vander Kloet and Dickinson 1999). Combined analyses of nuclear ribosomal ITS and chloroplast matK sequences clustered V. parvifolium with V. ovalifolium and indicated this clade to be directly ancestral the V. deliciosum clade (Powell and Kron 2002). The genetic differentiation observed in this study indicates greater diversity is present in V. parvifolium than previously reported in Northern California. The Shasta County and Sierra Ne- vada populations are more similar genetically to the congener V. deliciosum than they are to the coastal populations. Possible taxonomic revisions may involve expanding the definition of V. parvifolium to allow variation in berry color and high levels of genetic differentiation. Alternative- ly, a novel Vaccinium species or subspecies may be necessary to accurately reflect the morpholog- ical and genetic variation. Given the divergence of the Shasta County and Sierra Nevada populations from the typical V. parvifolium, the latter option may be more accurate. Additional morphological and ecologic study of these populations is warranted. Ecological and Conservation Implications Given the important role of Vaccinium shrubs in coniferous forests (Vila et al. 2004; Wender et al. 2004; Hanley 2005), an accurate understanding of the species diversity and distribution is necessary to understand the ecological role of this genus. The genetic differentiation observed among popu- lations of Vaccinium in northern California may reflect functional or adaptive divergence among geographic regions, in particular with regard to fruit color. Avian preference of berry color may vary temporally and with background foliage color (Burns and Dalen 2002; Honkavaara et al. 2004). In Rubus spectabilis Pursh, seed germina- tion of different color morphs varies with soil type, indicating fruit color may correlate with adaptive differentiation (Traveset and Willson 1998). In addition to berry color, differences in the extent of clonal growth patterns can affect the functional structure of populations. Clonal growth forms may allow genets to survive over a much longer timespan than individual ramets, and can significantly decrease effective population size (Persson and Gustavsson 2001). Observation- al and manipulative studies will be required to confirm if these differences constitute ecologically significant variation. While V. parvifolium is not a species of conservation concern, management plans should account for the distinct gene pools identified in 2012] these Vaccinium collections. The distinct mor- photype from Shasta County is of interest to USDA Forest Service management plans as it is consistent with growing evidence of a distinctive biodiversity in the Shasta Lake region, particu- larly if distinct subspecies or cryptic species are designated. Given the significant genetic differ- entiation among geographic groups (coastal, Shasta County, and Sierra Nevada), artificial movement of germplasm should be restricted to within and not among regions. These observa- tions are interesting as the Vaccinium berries are fleshy and palatable, and provide forage for numerous species in coniferous forests (Wender et al. 2004), though the dispersal ranges are likely small relative to the habitat range described for V. parvifolium (Rosatti 2003). ACKNOWLEDGMENTS The authors thank A.E.L. Colwell for floral descrip- tions from living and herbarium specimens. A.E.L. Colwell and G. Wallace provided helpful comments on the manuscript. J. Mello, R. Hernandez, R. Meyer, and R. 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WALDEN Department of Integrative Biology, University of California, Berkeley, CA 94720 gkwalden@berkeley.edu ROBERT PATTERSON Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132 ABSTRACT Nomenclature of subdivisions within Phacelia Juss. (Boraginaceae: Hydrophylloideae) reflects an update to the classification of the genus, based largely upon the structure offered in summary by Ferguson. We consider this proposed classification a continuation of efforts to better understand interrelationships within the genus and tribe Romanzoffieae Dumort., and anticipate future research offering insights into systematics of Phacelia. New names, changes in status, and combinations include the following: Phacelia sect. Baretiana Walden & R. Patt., sect. nov.; P. subsect. Bipinnatifidae (Small) Walden & R. Patt., stat. nov.; P. subsect. Cosmantha (Nolte ex A. de Candolle) Walden & R. Patt., stat. nov.; P. subsect. Cosmanthoides (A. Gray) Walden & R. Patt., stat. nov.; P. subsect. Dubiae (Small) Walden & R. Patt., stat. nov.; P. subsect. Eutoca (R. Br.) Walden & R. Patt., stat. nov.; P. sect. Glandulosae (Rydb.) Walden & R. Patt., stat. nov.; P. subsect. Humiles Walden & R. Patt., subsect. nov.; P. subsect. Lineares (Rydb.) Walden & R. Patt., stat. nov.; P. sect. Pachyphyllae Walden & R. Patt., sect. nov.; P. subg. Pulchellae (Rydb.) Walden & R. Patt., stat. nov.; P. sect. Ramosissimae (Rydb.) Walden & R. Patt., stat. nov.; P. subsect. Ranunculacea Walden & R. Patt., subsect. nov.; and P. subsect. Sericeae (Rydb.) Walden & R. Patt., stat. nov. Key Words: Boraginaceae, Hydrophyllaceae, Hydrophylloideae, Phacelia, Romanzoffieae. Philibert Commerson (variously Commerson, Commercon, or Commercon) and Jeanne Baret (variously Jean Baret, Jeanne Bare, Bonnefoi, or Bonnefoy) collected the first specimens of Pha- celia Juss. in the Straits of Magellan in the winter of 1767-1768 (Lamarck 1792). Commerson was the naturalist on the Bougainville expedition and called his own fascination with plants a “‘bota- nomania”’ (Bougainville 1771; Oliver and Elliot 1909). Baret, mistress of and field assistant to her “lover-master’’ Commerson, was disguised as a man for the majority of the lengthy sea journey and identified as a woman only when the expedition reached Tahiti (Dunmore 2002; Schie- binger 2003). Baret was the first woman to voyage around the world and was lauded as a skilled botanist (Bougainville 1771; Dunmore 2002; Schiebinger 2003; Ridley 2010; Tepe et al. 2012). Botanical collections from Brazil were lost on a return voyage across the Atlantic, and Commerson retained the remainder of his collec- tions, including those from the Straits of Magel- lan, until his death (Godley 1965). Commerson willed his herbarium to the Muséum national d’Histoire naturelle in Paris, and from these collections de Jussieu described Phacelia as a genus; P. secunda was later described and designated the type species (Jussieu 1789; Gmelin 1791; Laissus 1978). Taxonomic confusion began at the very formation of Phacelia, as researchers proposed different classifications and alliances for the variation observed from the Commerson collection. “L’ Hydrophylle de Magellan” (Hydro- Phyllum magellanicum Lam.) was also published from “‘Vherbier de Commerson,” (Lamarck 1792; Coville 1893; Deginani 1982). Although the informal ‘*Magellanicae’ has been generally adopted for the group of perennial species (e.g., Phacelia magellanica polyploid complex, Phacelia “species group Magellanicae’’), the type species of the genus is also included within the subdivision and so the epithet for the subsection is Phacelia (Heckard 1960; Constance and Chuang 1982). Gray (1875) combined the genera of A. de Candolle (1845) within Phacelia as subgenera, establishing a structure within the genus that has remained largely unchanged. In the first edition of the Manual of Botany, Gray (1848) noted subgenera and sections with the same mark, 8. Clarification came in the second edition, with subgenera and sections both noted with 8, but subgenera in all capitals (Gray 1856). In the Synoptical Flora, Gray (1878) defined the use of symbols and rank: ““The characters of sections of genera, when of comparatively high rank, are designated by the sectional mark (8) and printed in the larger type; and those of first importance, such as may be termed subgenera, are distin- guished by having a substantive name. Subsec- tions, and also primary divisions when of low 212 rank, are in small type.” These clarifications with discussion illustrated the need for hierarchical subdivisions in Phacelia and demonstrated ac- knowledgment of the intended use in the field (Candolle 1867; Brizicky 1968, 1969). Gray’s taxonomy developed with each iteration of published subdivisions within Phacelia, although adoption and application were not standardized. Nomenclature provided structure and guidance as contemporaries of Gray followed or chal- lenged his revisions and additions to the flora of North America. The taxonomy of Phacelia was complicated when Bentham and Hooker (1876) inadvertently recombined taxa by citing the subgenera of Gray (1875) but mistakenly trans- lated (8) at sectional status, considered a biblio- graphic error of citation, but a valid publication with priority (Brizicky 1968, 1969; Moore 2001; McNeill et al. 2006). Gray (1878) combined subgenera at sectional rank, citing himself, in his next publication, making these sectional names later isonyms, which may be disregarded (Bentham and Hooker 1876; McNeill et al. 2006). Later authors, such as Brand (1913), followed and perpetuated sectional combinations of Gray, which resulted in common use of names with no nomenclatural standing. We correct the sectional nomenclature of Phacelia in the taxonomic treatment. Brand’s (1913) classification of sections, infor- mal “‘conspectus varietatum,”’ and informal “‘sys- tema speciei’ has been the basis and inspiration for revisional taxonomic work in the genus, due in part to the large scale of his revision within Hydrophyllaceae. Characters of importance used to separate subdivisions in Phacelia were ovule and seed number, seed shape and _ surface morphology, and corolla scales (known as corolla squamae, corolla plicae, interstaminal scales, appendages, or lamellae) (Candolle 1845; Gray 1875, 1878; Munz 1935; Constance 1949; Gillett 1968; Hoffmann 1999). Rydberg (1917) limited his flora to species occurring within geographical bounds of the Rocky Mountains and organized Phacelia into seven unranked, but validly published, named subdivisions. Following a long-term study of chromosome numbers, Constance (1963) pro- posed a classification of three subgenera and ten informal “‘species groups.” Investigations of pollen surface morphology, trichomes, and seed surface morphology resulted in expanded taxo- nomic characters for the genus (Atwood 1975; Halse 1979; Constance and Chuang 1982; Di Fulvio and Dottori 1995). Ferguson (1998) offered an update to Constance’s classification, and although not a formal taxonomic revision, the synopsis of subdivisions in Phacelia, recog- nized to contain three subgenera, five sections, and six informal species groups, and broad sampling of genera within Hydrophyllaceae, MADRONO [Vol. 59 offered a structure for future taxonomic direc- tions in the genus. The monospecific Phacelia subg. Howellanthus Constance (1953) has since been removed to Howellanthus (Constance) Walden & R. Patt., and although we assigned the genus to tribe Phacelieae Benth. ex A. Gray (1875), Reveal’s Indices Supragenericorum Nomi- num Plantarum Vascularium (2012) identified Romanzoffieae Dumort. (Dumortier 1829) as having priority at rank tribe, and we correct our error here (Walden and Patterson 2010). Phacelia subg. Cosmanthus (Nolte ex A. de Candolle) A. Gray contains 20 species, P. subg. Phacelia encompasses the remaining 187 species across five sections, and 50 species were not assigned to a species group within P. sect. Phacelia (Ferguson 1998). By reviewing proto- logues, revisions, and molecular studies, we clarify the status of those unassigned taxa within this proposed classification. With additional species descriptions and systematic studies in Phacelia since Ferguson (1998), there is a need to formally recognize subdivisions in the genus, combine some previously separate subdivisions (P. subg. Phacelia and P. subg. Eutoca), provide names for other subdivisions (see Appendix | for outline), and note traditional taxonomic charac- ters that are useful for classification and identi- fication in a key to subdivisions. PHACELIA SUBG. PHACELIA Howell published revisions on two groups of annual species with entire-margined leaves from P. sect. Euphacelia (Howell 1943b) and P. sect. Eutoca (Howell 1945), later combined as the informal “‘species group Humiles’’ by Constance (1963). Lee (1986) examined the systematics of “species group Humiles’’ using corolla venation patterns and identified five morphological groups. Molecular studies of Phacelia support a sister relationship between the annual “‘species group Humiles’ and perennial “species group Magellanicae,”’ which we recognize formally here as P. subsect. Humiles and P. subsect. Phacelia within P. sect. Phacelia (Ferguson 1998; Gilbert et al. 2005; Walden 2010). It is clear that the nomenclature and taxonomic relationships of South America annual and perennial taxa should be reconsidered (Deginani 1982). Molecular studies have supported a clade consisting of three perennial species: P. hydro- phylloides Torr. ex A. Gray, P. procera A. Gray, and P. bolanderi A. Gray (Ferguson 1998; Gilbert et al. 2005; Walden 2010). These species have been traditionally included within Phacelia subg. Eutoca A. Gray, and were described together in one publication (Gray 1875). We treat this perennial group as P. sect. Baretiana within P. subg. Phacelia, named to honor the contributions of Baret to the botanical history of the genus, her 2012] long life, and long journey to public recognition (Walden 2010). “Species group Crenulatae” 1s an assemblage of 50 species, traditionally grouped by the morphological characteristics of trichomes stipi- tate-glandular with unicellular or multicellular heads, plants generally mephitic or malodorous, seeds cymbiform and excavated along one or both sides of a central adaxial ridge, seed surface reticulate-pitted and sometimes alveolate, and n = 11 (Voss 1937a, b; Atwood 1975; Garrison 2007; Walden 2010). Brand (1913) first grouped species within the informal “P. crenulata con- spectus varietatum,” ““P. glandulosa systema speciei,’ and ‘‘P. neo-mexicana systema speciei,” in his monograph of Hydrophyllaceae. Rydberg’s (1917) Phacelia [unranked] Glandulosae some- what encompassed the informal groupings of Brand (1913). Voss (1937a, b) revised the ‘“*Phacelia Crenulatae group,” an informal name that has stayed with the subdivision (Constance 1963; Atwood 1975). Taxa are distributed from Wyoming to México, with an amphitropical disjunction of three taxa in South America (P. artemisioides Griseb., P. pinnatifida Griseb. ex Wedd., and P. setigera Phil.) (Deginani 1982). Molecular studies have supported sampled “‘spe- cies group Crenulatae’ as monophyletic, al- though clearly not limited to a four-seeded capsule, and sister to a monophyletic “‘species group Tanacetifoliae’ [treated here as P. sect. Ramosissimae| (Gilbert et al. 2005; Garrison 2007; Hansen et al. 2009; Walden 2010). We treat Rydberg’s validly published name as the basionym for P. sect. Glandulosae within P. subg. Phacelia. When Gray (1875) established P. subg. Cos- manthus and P. subg. Cosmanthoides, he did so by splitting Cosmanthus Nolte ex A. de Candolle sect. Eucosmanthus, and assigning the majority of species to P. subg. Cosmanthoides. The typifica- tion of each subdivision has not been made explicit, and we provide lectotypification here. Orthographic changes in gender are required in subdivisions combined from Cosmanthus to Phacelia. In the preface to his Manual, Small (1933) wrote: ““Complex genera have been divided into more natural groups, both for convenience of study and also in order to make the genera, as far as possible, correspond to the great majority of groups of species now recog- nized as genera by most present-day botanists.” The unranked subdivisions of Small (1933) are treated as the basionyms for P. subsect. Bipinna- tifidae and P. subsect. Dubiae. Constance (1949, 1950) authored revisions of P. subg. Cosmanthus, documenting the distribution of the group from northeastern United States into México and Guatemala. Gillett (1968, pg. 368) noted “‘a basis for deleting the subgenus Cosmanthus as a systematic group’; in its place he proposed five WALDEN AND PATTERSON: PHACELIA SUBDIVISIONS 213 informal groups from biosystematic studies (Gillett 1964, 1965a, b, 1968). Molecular studies have maintained the monophyly of this subdivi- sion, although with limited sampling, supported as sister to Gillett’s “species group Franklinii”’ (treated here as P. sect. Eutoca) and nested within P. subg. Phacelia (Ferguson 1998; Gilbert et al. 2005; Hansen et al. 2009; Walden 2010). We treat P. sect. Cosmanthus within P. subg. Phacelia. Gillett’s “‘P.. ranunculacea Group” has been further explored by Sewell and Vincent (2009), and by Glass and Levy (2011), and we treat this clade as P. subsect. Ranunculacea within P. sect. Cosmanthus. Gray (1875) included Eutoca R.Br. within Phacelia, justifying the merger by noting that Phacelia and Eutoca were ‘“‘polymorphous”’ (Gray 1875). Rydberg’s (1917) Phacelia [un- ranked] Lineares and Phacelia [unranked] Ser- iceae were investigated in a series of biosystematic studies by Gillett (1960a, b, 1961, 1962, 1963) as the informal “‘species group Franklinii.”’ We treat Rydberg’s taxa as P. subsect. Lineares and P. subsect. Sericeae within P. sect. Eutoca. PHACELIA SUBG. MICROGENETES Molecular studies have supported a monophy- letic clade formed of three species: P. pachyphylla A. Gray, P. calthifolia Brand, and P. neglecta M. E. Jones, sister to P. sect. Euglypta and P. sect. Miltitzia (Dempcy 1996; Ganong 2002; Gilbert et al. 2005). Gray (1883) noted in his description of P. pachyphylla, “‘A most peculiar species, to be placed at the end of the Microgenetes section.” Howell (1942) provided a diagnosis for the informal group within a key when forming the “compact triad among our desert phacelias,”’ and Gilbert et al. (2005) noted, “All analyses sup- ported Howell’s (1946) Phacelia pachyphylla complex as a distinct lineage.” We treat this group as P. sect. Pachyphyllae within P. subg. Microgenetes. PHACELIA SUBG. PULCHELLAE Rydberg’s (1917) Phacelia [unranked] Pulchel- lae, diagnosed within keys, contained ten species of annual or perennial plants of low habit, corollas open-campanulate to tubular, corollas with stamens included, reticulate-pitted seeds, leaves long-petiolate, and leaf margins entire to lobed. Although only implied, the sharing of similar characteristics of Phacelia [unranked] Pulchellae to Phacelia [unranked] Bicolores sug- gested a close relationship between the two subdivisions (Torrey 1871; Gray 1875; Rydberg 1917). In discussion, Howell (1943a) referred to the “‘section Pulchellae”’ of Rydberg, but it does not appear that was a formal publication at that rank. Howell’s (1943a) revision included 19 214 species of annual or perennial plants with tubular corollas, stamens included, styles shallowly bifid, leaf margins entire to shallowly lobed, capsules elliptic or oblong, and ovules more than four per ovary. Phacelia [unranked] Pulchellae, P. sect. Euglypta, P. sect. Miltitzia, and Romanzoffia Cham. have tricolpate pollen without pseudo- colpi; the remainder of sampled Phacelia share tricolpate-tripseudocolpate pollen (Ferguson 1998). Phacelia [unranked] Pulchellae, consisting of the species around which Howell (1943a) centered his revision, was supported to have basal placement within the genus in molecular studies, sister to P. subg. Phacelia (Ferguson 1998; Gilbert et al. 2005; Walden 2010). We treat Rydberg’s validly published name as the basio- nym for P. subg. Pulchellae. TAXONOMIC TREATMENT Phacelia Juss., Gen. Pl. 129. 1789. —Type (lectotype designated by J. F. Gmelin 1791): Phacelia secunda J. F. Gmel., Syst. Nat. ed. 13. 2:330..1791. Aldea Ruiz & Pavon, Fl. Peruv. Prodr. 16—17. 1794[1798]. —Type (lectotype designated by Ruiz & Pavon, 1799): Aldea pinnata Ruiz & Pavon, FI. Peruv. 2:8, pl. 114 [CXIV], fig. a. 1799 (refers to p. 19 of the protologue in error). Phacelia Juss. subg. Phacelia. Phacelia Juss. sect. Phacelia. Phacelia Juss. subsect. Phacelia. Phacelia [unranked] Heterophyllae Rydb., FI. Rocky Mts. 702. 1917. —Type: Phacelia heterophylla Pursh, Fl. Amer. Sept. 140. 1814. Included taxa: Phacelia argentea A. Nelson & J. F. Macbr., P. californica Cham., P. capitata Kruckeb., P. corymbosa Jeps., P. egena (Brand) J. T. Howell, P. hastata Douglas ex Lehm., P. hastata var. hastata, P. hastata var. charlestonen- sis Cronquist, P. hastata var. compacta (Brand) Cronquist, P. heterophylla Pursh, P. heterophylla var. heterophylla, P. heterophylla var. virgata (Greene) Dorn, P. imbricata Greene, P. imbricata var. imbricata, P. imbricata var. bernardina (Greene) Walden & R. Patt., P. imbricata var. patula (Brand) Walden & R. Patt., P. /eptosepala Rydb., P. mutabilis Greene, P. nemoralis Greene, P. nemoralis var. nemoralis, P nemoralis var. oregonensis (Heckard) Walden & R. Patt., P. secunda J. F. Gmel., P. secunda var. secunda, P. secunda var. pinnata (Vahl) Deginani. Phacelia Juss. subsect. Humiles Walden & R. Patt., subsect. nov. —Type: Phacelia humilis Torr. & A. Gray, in War Department (U.S.), Pacif. Railr. Rep. 2:122. 1855. Plants annual; herbage mephitic or unscented. Stems decumbent to ascending to erect, sometimes MADRONO [Vol. 59 wiry, simple, or branched; hirtellous to hispidu- lous to hirsute, eglandular or glandular, glands colorless- to amber- or dark-tipped. Leaves rosulate, or opposite proximally and alternate distally, or alternate and cauline; petiolate or subsessile; blade linear to lanceolate to oblong or narrowly ovate, simple and margins entire, or margins shallowly pinnatifid or toothed or shal- lowly lobed with 1-4 pairs, lobe margins entire, bases cuneate or attenuate, hirtellous to hispidu- lous to hirsute, eglandular or glandular. Inflores- cence unit a cyme, secund or racemose, usually solitary, or sometimes geminate, ascending to erect, one- or two-ranked, extending or not beyond vegetation. Flowers pedicellate, pedicels short or long, spreading to ascending, or arcuate in fruit; calyx slightly accrescent or strongly accrescent in fruit, lobes equal or unequal, linear to oblanceolate to spatulate to obovate, some- times foliaceous, hirsute and glandular, margins ciliate; corollas deciduous or marcescent, cam- panulate to open-campanulate to subrotate, white or blue or lavender or purple, lamina sometimes with translucent areas, glabrous adaxially, puber- ulent to sparsely hirsute abaxially, lobe margins entire or erose; nectary gland usually absent, rarely present (P. doug/asii); corolla scales usually present, sometimes absent, linear to lanceolate to ovate, adjacent scale edges divergent or not across base of filament, glabrous or ciliate; stamens included or exsert, filaments equal or unequal in length, glabrous or papillate, anthers bronze; style included or exsert, branched 1/2 length to branched nearly to base, hirsute proximally; ovules 2—12 per placenta. Fruits ellipsoid to ovoid and compressed along sutures, or globose and plump, hirsute and sparsely glandular. Seeds 2— 20, brown, oblong or ovoid, rounded or truncate at ends, sometimes angled, surface finely or coarsely reticulate-pitted, reticulations rarely in transverse striations. n = 7, 8, 9, 10, 11. Included taxa: Phacelia austromontana J. T. Howell, P. brachyantha Benth., P. breweri A. Gray, P. congdonii Greene, P. curvipes Torr. ex S. Watson, P. davidsonii A. Gray, P. divaricata (Benth.) A. Gray, P. douglasii (Benth.) Torr., P. eisenii Brandegee, P. exilis (A. Gray) G. J. Lee, P. greenei J. T. Howell, P. grisea A. Gray, P. humilis Torr. & A. Gray, P. humilis var. humilis, P. humilis var. dudleyi J. T. Howell, P. inconspicua Greene, P. insularis Munz, P. insularis var. insularis, P. insularis var. continentis J. T. Howell, P. leonis J. T. Howell, P. marcescens Eastw. ex J. F. Macbr., P. minutissima L. F. Hend., P. mohavensis A. Gray, P. novenmillensis Munz, P. orogenes Brand, P. peckii J. T. Howell, P. phacelioides A. Gray, P. pringlei A. Gray, P.:purpusii Brandegee, P. quickii J.T. Howell, P. racemosa (Kellogg) Brandegee, P. stebbinsii Constance & Heckard, P. stellaris Brand, P. vallicola Congdon ex Brand, P. verna J. T. Howell. 2012] Phacelia Juss. sect. Baretiana Walden & R. Patt., sect. nov. —Type: Phacelia hydrophylloides Torr. ex A. Gray, Proc. Amer. Acad. Arts 7(2):400. 1868. Plants perennial; herbage malodorous. Stems decumbent to ascending to erect; usually hirtel- lous to hirsute, glandular, glands colorless- to amber- to dark-tipped, sometimes glabrate prox- imally. Leaves petiolate; blade oblong to ovate to subrhombic, simple or pinnatifid to lyrate or pinnate with 1-3 pairs of leaflets at base, leaflets oblong to ovate, bases attenuate or truncate to subcordate, margins usually incised or serrate or dentate, rarely subentire, faces usually hirtellous to hirsute, sometimes glabrate, margins some- times hispid-ciliate, glandular. Inflorescence unit a cyme, paniculate or capitate, solitary or in 2-3 clusters. Flowers pedicellate, pedicels short or long, straight in fruit; calyx slightly accrescent in fruit, lobes equal, linear to oblanceolate to oblong to narrowly spatulate, hirsute and glan- dular, margins hispid-ciliate, tips spreading; corollas deciduous, rotate to open-campanulate, white to cream to green-white or pale blue to lavender, throat sometimes fading brown in age, glabrous adaxially, puberulent abaxially, lobe margins entire or erose, spreading or revolute; nectary gland absent; corolla scales present, oblong, adjacent scale edges divergent across bases of filaments, scale edges sometimes adnate and forming a narrow sac, glabrous; stamens slightly exsert to exsert, filaments equal or slightly unequal, glabrous or hirsute, anthers blue or purple-brown or white; style included or exsert, branched 1/2 to 3/4 length, hirsute proximally; ovules 3—30 per placenta. Fruits plumply ovoid to subglobose, apiculate, hirsute. Seeds 3—60, brown or black, oblong to ellipsoid to irregularly cylindrical, angled, acute at both ends, adaxial surface sometimes with shallow keel, surface shallowly foveolate or finely scrobiculate, reticu- late-pitted. n = 11. Included taxa: Phacelia bolanderi A. Gray, P. hydrophylloides Torr. ex A. Gray, P. procera A. Gray. Phacelia Juss. sect. Cosmantha (Nolte ex A. de Candolle) Benth. & Hook.f., Gen. Pl. 2:828. 1876. Cosmanthus Nolte ex A. de Candolle, Prodr. 9:296. 1845. Phacelia subg. Cosmanthus (Nolte ex A. de Candolle) A. Gray, Proc. Amer. Acad. Arts 10:320. 1875 (as 8 2. COSMANTHUS). —Type (lectotype designat- ed here): Phacelia fimbriata Michx., Fl. Bor.- Amer. 1:134. 1803. Phacelia subsect. Cosmantha (Nolte ex A. de Candolle) Walden & R. Patt., stat. nov. Phacelia [unranked] Fimbriatae Small, Man. S.E. FI. 1097. 1933. —Type (lectotype designated here): WALDEN AND PATTERSON: PHACELIA SUBDIVISIONS 2S Phacelia fimbriata Michx., Fl. Bor.-Amer. 1:134. 1803. Included taxa: Phacelia fimbriata Michx., P. purshii Buckley Phacelia Juss. subsect. Bipinnatifidae (Small) Walden & R. Patt., stat. nov. Phacelia [un- ranked] Bipinnatifidae Small, Man. S.E. FI. 1097. 1933. —Type: Phacelia bipinnatifida Michx., Fl. Bor.-Amer. 1:134. 1803. Included taxon: Phacelia bipinnatifida Michx. Phacelia Juss. subsect. Cosmanthoides (A. Gray) Walden & R. Patt., stat. nov. Phacelia subg. Cosmanthoides A. Gray, Proc. Amer. Acad. Arts 10:320. 1875 (as 8 3. COSMANTHOIDES). Phacelia sect. Cosmanthoides (A. Gray) Benth. & Hook.f., Gen. Pl. 2:828. 1876. —Type (lectotype designated here): Phacelia platycarpa (Cav.) Spreng., Syst. Veg. (ed. 16) [Sprengel] 1:584. 1824 [1825]. Included taxa: Phacelia altotonga B. L. Turner, P. austrotexana (J. A. Moyer) B. L. Turner, P. carmenensis B. L. Turner, P. gilioides Brand, P. glabra Nutt., P. hirsuta Nutt., P. /axa Small, P. neffii B. L. Turner, P. patuliflora A. Gray, P. platycarpa (Cav.) Spreng., P. platycarpa var. platycarpa, P. platycarpa var. bursifolia (Willd. ex Roem. & Schult.) Constance, P. platycarpa var. madrensis (Greenm.) Constance, P. pulcher- rima Constance, P. strictiflora (Engelm. & A. Gray) A. Gray, P. strictiflora var. strictiflora, P. strictiflora var. connexa Constance, P. strictiflora var. lundelliana Constance, P. strictiflora var. robbinsii Constance, P. teucriifolia 1. M. Johnst., P. zaragozana B. L. Turner. Phacelia Juss. subsect. Dubiae (Small) Walden & R. Patt., stat. nov. Phacelia [unranked] Dubiae Small, Man. S.E. Fl. 1097. 1933. —Type: Phacelia dubia (L.) Trel. & Small, Rep. (Annual) Arkansas Geol. Surv. (for 1888). 4:205. 1891. Included taxa: Phacelia dubia (L.) Trel. & Small, P. dubia var. dubia, P. dubia var. georgiana McVaugh, P. dubia var. interior Fernald, P. maculata Wood. Phacelia Juss. subsect. Ranunculacea Walden & R. Patt., subsect. nov. —Type: Phacelia ranun- culacea (Nutt.) Constance, Rhodora 42:39. 1940. Plants annual, (5)10—-25 cm; herbage unscent- ed. Stems prostrate to erect; hirsute and glandu- lar, glands colorless-tipped. Leaves blade oblong to ovate, pinnatifid or pinnate with 2—6 pairs of leaflets, lobes oblong to round, bases cuneate, margins entire or toothed, hirsute and glandular. 216 Inflorescence unit a cyme, secund, 1—6 flowers. Flowers pedicellate, pedicels short or long, reflexed to pendent in fruit; calyx slightly accrescent in fruit, lobes unequal, linear to lanceolate, hirsute and sparsely glandular; corol- las tubular-campanulate, pale violet or lavender, lobe margins entire; stamens included, filaments equal, glabrous; style included, branched 1/3 to 2/ 3 length, glabrous; ovules 2 per placenta. Fruits depressed globose, hirsute. Seeds 2-4, globose to ovoid, surface finely reticulate-pitted. n = 6, 14. Included taxa: Phacelia covillei S. Watson ex A. Gray, P. ranunculacea (Nutt.) Constance. Phacelia Juss. sect. Eutoca (R. Br.) Benth. & Hook. f., Gen. Pl. 2: 828. 1876. Eutoca R. Br., Narr. Journey Polar Sea. 764-765, tab. 27. 1823. Eutoca R. Br. sect. Ortheutoca A. de Candolle, Prodr. 9:296. 1845. Phacelia subg. Eutoca (R. Br.) A.Gray, Proc. Amer. Acad. Arts 10:322. 1875 (as 8 6. EUTOCA). —Type: Phacelia franklinii (R.Br.) A.Gray, Manual ed. 2:329. 1856. Phacelia subsect. Eutoca (R.Br.) Walden & R. Patt., stat. nov. —Type: Phacelia franklinii (R.Br.) A.Gray, Manual ed. 2:329. 1856. Included taxon: Phacelia franklinii (R. Br.) A. Gray. Phacelia Juss. subsect. Lineares (Rydb.) Walden & R. Patt., stat. nov. Phacelia [unranked] Lineares Rydb., Fl. Rocky Mts. 702. 1917. — Type: Phacelia linearis (Pursh) Holz., Contr. U.S. Natl. Herb. 3:242. 1895. Included taxon: Phacelia linearis (Pursh) Holz. Phacelia Juss. subsect. Sericeae (Rydb.) Walden & R. Patt., stat. nov. Phacelia [unranked] Sericeae Rydb., Fl. Rocky Mts. 702. 1917. — Type: Phacelia sericea (Graham) A. Gray, Amer. J. Sci. Arts, ser. 2. 34(101):254. 1862. Included taxa: Phacelia idahoensis L. F. Hend., P. lenta Piper, P. lyallii Rydb., P. mollis J. F. Macbr., P. sericea (Graham) A. Gray, P. sericea var. sericea, P. sericea var. ciliosa Rydb. Phacelia Juss. sect. Glandulosae (Rydb.) Walden & R. Patt., stat. nov. Phacelia [unranked] Glandulosae Rydb., Fl. Rocky Mts. 702. 1917. —-Type: Phacelia glandulosa Nutt., J. Acad. Nat. Sci. Philadelphia, ser. 2, 1:160. 1847. Included taxa: Phacelia alba Rydb., P. amabilis Constance, P. anelsonii J.F.Macbr., P. argylensis N. D. Atwood & S. L. Welsh, P. argillacea N. D. Atwood, P. arizonica A. Gray, P. artemisioides Griseb., P. bakeri (Brand) J. F. Macbr., P. bombycina Wooton & Standl., P. cloudcroftensis MADRONO [Vol. 59 N. D. Atwood, P. coerulea Greene, P. congesta Hook., P. constancei N. D. Atwood, P, corrugata A. Nelson, P. cottamii N. D. Atwood, P. coulteri Greenm., P. crenulata Torr. ex S. Watson, P. crenulata var. crenulata, P. crenulata var. ambigua (M. E. Jones) J. F. Macbr., P. crenulata var. angustifolia N. D. Atwood, P. crenulata var. minutiflora (J. W. Voss ex Munz) Jeps., P. denticulata Osterh., P. formosula Osterh., P. furnissii N. D. Atwood, P. glandulosa Nutt., P. glandulosa var. glandulosa, P. glandulosa var. deserta Brand, P. gypsogenia 1. M. Johnst., P. higginsti N. D. Atwood, P. hintoniorum B. L. Turner, P. howelliana N. D. Atwood, P. hughesii N. D. Atwood, P. infundibuliformis Torr., P. infundibuliformis var. infundibuliformis, P. infun- dibuliformis var. phanerandra 1. M. Johnst., P. integrifolia Torr., P. integrifolia var. integrifolia, P. integrifolia var. texana (J. W. Voss) N. D. Atwood, P. mammillarensis N. D. Atwood, P. marshall-johnstonii N. D. Atwood & Pinkava, P. marshall-johnstonii var. marshall-johnstonii, P. marshall-johnstonii var. deliciasana B. L. Turner, P. neomexicana Yhurb. ex Torr., P. orbicularis Rydb., P. pallida 1. M. Johnst., P. palmeri Torr. ex S. Watson, P. pedicellata A. Gray, P. petrosa N. D. Atwood, F. J. Sm., & T. A. Knight, P. pinkavae N. D. Atwood, P. pinnatifida Griseb. ex Wedd., P. popei Torr. & A. Gray, P, potosina B. L. Turner, P. rafaelensis N. D. Atwood, P. robusta (J. F. Macbr.) I. M. Johnst., P. rupestris Greene, P. sanzinii Hicken, P. scariosa Brande- gee, P. serrata J. W. Voss, P. setigera Phil., P. setigera var. setigera, P. setigera var. humahua- quense Deginani, P. sinuata Phil., P. sivinskii N. D. Atwood, P. J. Knight, & Lowrey, P. sonoitensis S. P. McLaughlin, P. splendens Eastw., P. utahensis J. W. Voss, P. vossii N. D. Atwood, P. welshii N. D. Atwood. Phacelia Juss. sect. Gymnobytha (A. de Candolle) Benth. & Hook.f., Gen. Pl. 2:828. 1876. Cosmanthus Nolte ex A. de Candolle sect. Gymnobythus A. de Candolle, Prodr. 9:296. 1845. Phacelia subg. Gymnobythus (A. de Candolle) A.Gray, Proc. Amer. Acad. Arts 10:321. 1875 (as 8 4. GYMNOBYTHUS). — Type: Phacelia viscida (Benth.) Torr., Rep. U.S. Mex. Bound., Bot. (Emory) 143. 1859. Included taxa: Phacelia grandiflora (Benth.) A. Gray, P. viscida (Benth.) Torr., P. viscida var. viscida, P. viscida var. albiflora (Nutt.) A. Gray. Phacelia Juss. sect. Ramosissimae (Rydb.) Walden & R. Patt., stat. nov. Phacelia [unranked] Ramosissimae Rydb., Fl. Rocky Mts., 702. 1917. —Type: Phacelia ramosissina Douglas ex Lehm., Nov. Stirp. Pug. (Lehmann) 2:21. 1830. Included taxa: Phacelia cedrosensis Rose, P. cicutaria Greene, P. cicutaria var. cicutaria, 2012] P. cicutaria var. hispida (A. Gray) J. T. Howell, i sciliata Benth:., P. cimerea Eastw. ex. J. F. Macbr., P. cryptantha Greene, P. distans Benth.., P. floribunda Greene, P. gentryi Constance, P. hirtuosa A. Gray, P. hubbyi (J. F. Macbr.) L. M. Garrison, P. ixodes Kellogg, P. lyonii A. Gray, P. malvifolia Cham. P. malvifolia var. malvifolia, P. malvifolia var. loasifolia (Benth.) Brand, P. phyllomanica A. Gray, P. platyloba A. Gray, P. pauciflora S. Watson, P. ramosissima Douglas ex Lehm., P. ramosissima var. ramosissima, P. ramosissima var. austrolitoralis Munz, P. ramo- sissima var. eremophila (Greene) J. F. Macbr., P. ramosissima var. latifolia (Torr.) Cronquist, P. ramosissima var. montereyensis Munz, P. rattanti A. Gray, P. tanacetifolia Benth., P. thermalis Greene, P. umbrosa Greene, P. vallis-mortae J. W. Voss. Phacelia Juss. sect. Whitlavia (Harv.) Benth. & Hook.f., Gen. Pl. 2:828. 1876. Whitlavia Harv., London J. Bot. 5:311-312, pl. Il. 1846. Phacelia subg. Whitlavia (Harv.) A. Gray, Proc. Amer. Acad. Arts 10:321. 1875 (as 8 5. WHITLAVIA). —Type: Phacelia minor (Harv.) Thell. ex F. Zimm., Ber. Bayer. Bot. Ges. 14:79. 1914. Phacelia subsect. Whitlaviae (Harv.) G. W. Gillett, Univ. Calif. Publ. Bot. 28:60. 1955. — Type: Phacelia minor (Harv.) Thell. ex F. Zimm., Ber. Bayer. Bot. Ges. 14:79. 1914. Included taxa: Phacelia minor (Harv.) Thell. ex F.Zimm., P. parryi Torr. Phacelia Juss. subsect. Campanulariae G. W. Gillett, Univ. Calif. Publ. Bot. 28:62. 1955. — Type: Phacelia campanularia A. Gray, Syn. FI. N. Amer. 2(1):164. 1878. Included taxa: Phacelia campanularia A. Gray, P. campanularia var. campanularia, P. campanu- laria var. vasiformis (G. W. Gillett) Walden & R. Patt., P. /ongipes Torr. ex A. Gray, P. nashiana Jeps. Phacelia Juss. subg. Microgenetes (A. de Can- dolle) A. Gray, Proc. Amer. Acad. Arts 10:326. 1875 (as 8 7. MICROGENETES). Microge- netes A. de Candolle, Prodr. 9: 292-293. 1845. —Type (lectotype designated by J. T. Howell 1946): Phacelia cumingii (Benth.) A. Gray, Syn. Fl. N. Amer. 2(1):169. 1878. Phacelia Juss. sect. Euglypta S. Watson, Botany (Fortieth Parallel) 254. 1871. Phacelia sect. Microgenetes (A. de Candolle) Benth. & Hook.f., Gen. Pl. 2:828. 1876. —Type (lecto- type designated by J. T. Howell 1946): Phacelia cumingii (Benth.) A. Gray, Syn. Fl. N. Amer. 2(1):169. 1878. WALDEN AND PATTERSON: PHACELIA SUBDIVISIONS 217 Phacelia [unranked] Bicolores Rydb., Fl. Rocky Mts. 702. 1917.—Type: Phacelia bicolor Torr. ex S.Watson, Botany (Fortieth Parallel) 255. 1871. Included taxa: Phacelia affinis A. Gray, P. bicolor Torr. ex S. Watson, P. brachyloba (Benth.) A. Gray, P. cephalotes A. Gray, P. cumingii (Benth.) A. Gray, P. fremontii Torr., P. glandu- lifera Piper, P. gymnoclada Torr. ex S. Watson, P. ivesiana Torr., P. leibergii Brand, P. nana Wedd. Phacelia Juss. sect. Miltitzia (A. de Candolle) J. T. Howell, Leafl. W. Bot. 4:15. 1944. Miltitzia A. de Candolle, Prodr. 9:296. 1845. Emme- nanthe Benth. subg. Miltitzia (A. de Candolle) A. Gray, War Department (U.S.), Pacif. Railr. Rep. 1854-5, 6:84-85. 1857 (Gray wrote, “‘It will be seen that I incline to the latter view; but should retain Mil/titzia as a subgenus.’’). — Type: Phacelia lutea (Hook. & Arnott) J. T. Howell, Leafl. W. Bot. 4:15. 1944. Included taxa: Phacelia adenophora J. T. Howell, P. glaberrima (Torr. ex S. Watson) J. T. Howell, P. inundata J. T. Howell, P. inyvoensis (J. F. Macbr.) J. T. Howell, P. /utea (Hook. & Arnott) J. T. Howell, P. /utea var. lutea, P. lutea var. calva Cronquist, P. /utea var. mackenzieorum J. W. Grimes & P. L. Packard, P. lutea var. purpurascens J.T. Howell, P. monoensis Halse, P. salina (A. Nelson) J. T. Howell, P. scopulina (A. Nelson) J. T. Howell, P. submutica J. T. Howell, P. tetramera J. T. Howell. Phacelia Juss. sect. Pachyphyllae Walden & R. Patt., sect. nov. —Type: Phacelia pachyphylla A. Gray, Proc. Amer. Acad. Arts 19:88. 1883. Plants annual, 3—35 cm, herbage mephitic. Stems erect, hirsute and glandular, glands colorless- to amber- to black-tipped. Leaves rosulate and long-petiolate proximally, reduced and subsessile distally, petioles stout, proximal petioles usually channeled; blade broadly ovate or reniform or round, simple, bases truncate or cordate, margins undulate to shallowly lobed or crenate or serrulate, thick and succulent to coriaceous, hirtellous and glandular adaxially, glabrate and glandular abaxially, veins im- pressed adaxially. Inflorescence unit a cyme, secund, erect, solitary or in 2-3 clusters, compact and not elongate proximally, extending or not beyond vegetation. Flowers pedicellate, pedicels short, spreading to arcuate proximally in fruit; calyx slightly accrescent in fruit, lobes equal, oblong to oblanceolate, pubescent to hirsute and glandular; corollas deciduous, fun- nelform-campanulate or open-campanulate or subrotate, tube white to purple, lobes white to purple, glabrous adaxially, puberulent abaxi- ally, lobe margins entire; nectary gland absent; corolla scales reduced and narrow, adjacent scale 218 edges sometimes divergent and adnate across base of filament, hirsute or ciliate; stamens included, filaments subequal or unequal, glabrous or puberulent, anthers yellow or violet; style includ- ed, branched 1/2 to 3/4 length, hirtellous and glandular; ovules 50—120. Fruits plumply ovoid to globose, prominently sulcate, puberulent and glandular. Seeds dark brown, 30—120, ellipsoid to ovoid, angular, surface transversely corrugat- ed, corrugations 4-8. n = 11, 12. Included taxa: Phacelia calthifolia Brand, P. neglecta M. E. Jones, P. pachyphylla A. Gray. Phacelia Juss. subg. Pulchellae (Rydb.) Walden & R. Patt., stat. nov. Phacelia [unranked] Pulchel- lae Rydb., Fl. Rocky Mts. 702. 1917. —Type: Phacelia pulchella A.Gray, Proc. Amer. Acad. Arts 10:326. 1875. MADRONO [Vol. 59 Included taxa: Phacelia barnebyana J. T. Howell, P. beatleyae Reveal & Constance, P. cookei Constance & Heckard, P. cronquistiana S. L. Welsh, P. demissa A. Gray, P. demissa var. demissa, P. demissa var. heterotricha J. T. Howell, P. demissa var. minor N. D. Atwood, P. filiae N. D. Atwood, F. J. Sm., & T. A. Knight, P. filiformis Brand, P. geraniifolia Brand, P. glecho- mifolia A. Gray, P. incana Brand, P. indecora J. T. Howell, P. keckii Munz & I. M. Johnst., P. laxiflora J. T. Howell, P. lemmonii A. Gray, P. mustelina Coville, P. parishii A. Gray, P. peirsoniana J. T. Howell, P. perityloides Coville, P. pulchella A. Gray, P. pulchella var. pulchella, P. pulchella var. gooddingii (Brand) J. T. Howell, P. rotundifolia Torr. ex S. Watson, P. sabulonum (J. T. Howell) N. D. Atwood, P. saxicola A. Gray, P. suaveolens Greene. KEY TO SUBDIVISIONS IN PHACELIA Plants 3—35 cm; corolla tubular to campanulate; corolla scales reduced and narrow, or absent; stamens included, subequal to unequal in length; style shallowly 2-lobed to branched 1/2 length; capsule costate and longitudinally sulcate 2. Annuals or perennials; seed compressed or angled, surface reticulate-pitted or foveolate........... Sait, Omen atk oe Be ur A, gues, oe ey eh eee aes Sen aliens eee, aoe Rawle Phacelia subg. Pulchellae 2’ Annuals; seeds terete and plump, surface transversely corrugated or transversely striate........... SS eee Soe OE ae ee B. Soe ae etek eat eee et ok ae ea Se ae Phacelia subg. Microgenetes 3. Leaf blades round, simple, bases cordate; corolla not yellow at base, tube white or purple; style branched 1/2 length; capsule globose, plump, exceeding calyx lobes; seeds 30-120 per CAS UlGie 2 ce ie rte oS agi eee te Re eee ieee) See Dense rena haan Oe Phacelia sect. Pachyphyllae 3’ Leaf blades oblong to ovate, usually pinnatifid to pinnate to bipinnatifid, rarely simple, bases cuneate to truncate; corolla yellow at base, tube white or yellow or blue; style shallowly 2-lobed to branched 1/4 length; capsule oblong to ellipsoid to cylindric, not plump, not exceeding calyx lobes (except P. tetramera); seeds 4-30 per capsule 4. Stems prostrate to ascending; herbage neither mephitic nor malodorous; corolla tardily deciduous or marcescent in fruit, lobes white or yellow to yellow-purple. .Phacelia sect. Miltitzia 4’ Stems ascending to erect; herbage usually mephitic or malodorous; corolla readily deciduous in fruit, lobes white or pink or purple or blue ................... Phacelia sect. Euglypta Plants 5—200 cm; corolla tubular or campanulate or rotate; corolla scales present in pairs along lateral veins of lamina (10 total in 5 pairs), reduced, or absent; stamens usually exsert, usually equal in length; style branched 1/4 length to parted nearly to base; capsule ecostate and shallowly longitudinally SUUCAIC Sse pracy ee Oe oe Aide, Pegs oe anette ren Pe opium eh ence ah Oe a) Eon Phacelia subg. Phacelia 5. Annuals; corolla scales absent; seed surface shallowly reticulate-pitted and foveolate 6. Staminal appendages at base of filaments absent; seeds 40-200 per capsule, ovules 50—120 per PMACCICA eS eee eee eet Sh, See yosiet rae ects co ae Oe Gio Rae eae Phacelia sect. Gymnobytha 6’ Staminal appendages at base of filaments present; seeds 10—90 per capsule, ovules 5-45 per POMACENNGA 5 cack ert Begs inen Sag ae te, a te, eae os eee eaters, ae Phacelia sect. Whitlavia 7. Corolla usually purple, rarely white, corolla markings opposite corolla lobes or none; staminal appendages hairy; seeds 1-1.5mm.................. Phacelia subsect. Whitlaviae 7’ Corolla white or pink to pale violet or bright blue, corolla markings opposite corolla sinuses or none; staminal appendages glabrous; seeds 1.5—-2 mm...... Phacelia subsect. Campanulariae 5’ Annuals, biennials, or perennials; corolla scales usually present, wholly or partially adnate to corolla tube along one or both scale edges, sometimes reduced or absent; seed surface reticulate-pitted, alveolate, or foveolate 8. Nectary gland present on lamina midvein, sometimes reduced to minute ridge; corolla scale distal edges bordering or overlapping midvein, not divergent proximally across base of filament 9. Corolla scales wholly adnate along one edge to lateral vein, distal free edge overlapping midvein or nectary gland; seeds 6—60 per capsule; western North America (usually west of Great Plains, except some P. franklinii populations)................. Phacelia sect. Eutoca 10. Perennials; nectary gland surface hairy; stamens exsert, filaments glabrous or SlADIAIC ag ts ie hen, be a eds a OS Oa ner Phacelia subsect. Sericeae 10’ Annuals or biennials; nectary gland surface glabrous; stamens included or equal to corolla, filaments hairy 11. Annuals; stamens included, filaments stipitate-glandular; seeds 6-15 per capsule, surtace coarsely reticulate-pitted | ..544 44. 24.5 > oe anes | Phacelia subsect. Lineares 2012] Q’ WALDEN AND PATTERSON: PHACELIA SUBDIVISIONS 219 11’ Biennials; stamens equal to corolla, filaments eglandular; seeds 40—60 per capsule, SuULiace finely Tretichlate-pitled <2 20s e 4n.4.5 as Deen oo oe % Phacelia subsect. Eutoca Corolla scales usually wholly adnate along one or both edges to lateral vein, free edge bordering and not overlapping midvein or nectary gland, sometimes each reduced to a ridge; seeds 2-20 per capsule; eastern North America (east of Great Plains), México, and GSeUWAte OA win 2.08 e. te hase woth & ye 2 araputn Made) Rana se Gp nie eeee oe eee Phacelia sect. Cosmantha 12. Annual or biennial herbs; seeds 2-4 per capsule, ovules 2 per placenta; seed globose or ovoid, not angled 13. Corolla tubular-campanulate; stamens included, filaments glabrous; seed adaxial keel USMY om: oS ok Anat, cain ap enema ens, SLMem nee DON Mit nce enh Phacelia subsect. Ranunculacea 13’ Corolla open-campanulate to rotate-campanulate; stamens equal to corolla or exsert, filaments hairy; seed adaxial keel present. 14. Biennial herbs; corolla lobe margins entire or erose; stamens exsert; seed surface coarsely reticulate-pitted, excavated along one side of adaxial IG lis, Seen, ee eee case Cae ook ee ate onset ee a Phacelia subsect. Bipinnatifidae 14’ Annual herbs; corolla lobe margins fimbriate; stamens equal to corolla; seed surface finely reticulate-pitted, not excavated alongside adaxial keel.......... Ae Fi See, in cae ea np RN a say Ace a gees Phacelia subsect. Cosmantha 12’ Annual or perennial herbs; seeds usually 5—20 per capsule, ovules usually 4—14 per placenta, rarely ovules 2 per placenta and seeds (2—3)4 per capsule (P. zaragozana); seed ovoid or ellipsoid, angled 15. Annual herbs; plants usually east of Mississippi River (not of México or TERESINA oe are, ase ree Se Ho eh aes Stet ey ee Ste ahs: Meow es Qed LE Phacelia subsect. Dubiae 15’ Annual or perennial herbs; plants usually west of Mississippi River (except some populations of P. strictiflora var. lundelliana, also of México, Guatemala) ......... Pes get ede Be Beek, Set enctee Oe e eeneee og, cana! oe ea te ea te Phacelia subsect. Cosmanthoides 8’ Nectary gland usually absent on lamina midvein, rarely present (P. douglasii); corolla scales usually divergent proximally across base of filament, distal scale edges between filaments free or connate, sometimes reduced or absent 16. Seed cymbiform (shallowly cymbiform in P. sonoitensis and P. infundibuliformis), usually excavated along one or both sides of adaxial ridge, forming two longitudinal grooves, sometimes shallowly excavated (P. bakeri), surface reticulate-pitted or alveolate, adaxial ridge sometimes corrugated, seeds sometimes marginate, margins sometimes corrugated ......... ae or eg ee es ae ae ees Phacelia sect. Glandulosae 16’ Seed terete or carinate or angled, not cymbiform, sometimes shallowly excavated alongside adaxial ridge or keel, surface reticulate-pitted or foveolate or alveolate or scrobiculate or rugose, adaxial ridge or keel not corrugated, seeds rarely marginate, margins not corrugated 17. Leaves cauline, alternate, usually pinnatifid or pinnate or bipinnatifid, rarely simple (P. bolanderi, P. malvifolia, and P. rattanii), margins lobed or pinnatifid 18. Annuals or perennials; annuals with seeds 1-40 per capsule, perennials with seeds 1-4 per capsule; calyx usually strongly accrescent in fruit, lobes usually unequal........ Phacelia sect. Ramosissimae 18’ Perennials; seeds 3-60 per capsule; calyx slightly accrescent in fruit, lobes SUG earn hee G24 4 eed 2 eases 6 2 oe cee ee ee Phacelia sect. Baretiana 17’ Leaves in basal rosette or rosulate, or first pair opposite to subopposite, or leaves cauline and alternate, simple or shallowly lobed and margins entire, or pinnatifid to pinnate and leat letanareins Civiite: ty. ees eet ee Sete Gee yy See ene te aaa Phacelia sect. Phacelia 19. Perennials or biennials; stamens exsert; seeds (1—2)3—4 per capsule, ovules 2 per PNAC CTA es ce ek fees a es eee see cere nec Hen che eal Sete es Be wate ws ore Phacelia subsect. Phacelia 19’ Annuals; stamens included or exsert; seeds (1—2)3—20 per capsule, ovules 2—20 per placentas, «24.6444 5444 08% ACKNOWLEDGMENTS We thank Frank Cipriano, Dennis Desjardin, Ellen Dean, Bruce Baldwin, Diane Ferguson, Richard Olmstead, Leigh Johnson, Laura Garrison, Debra Hansen, John Dempcy, Deb Trock, Nancy Morin, Ron Hartman, Jim Linnberg, and Trigger (service dog of GKW). We are grateful to archivists, librarians, and curators at many institutions for access to books, databases, specimens, and loans supporting this re- search. This represents, in part, a master’s thesis by GKW (2010) submitted to SFSU, and is in support of the FNANM treatment of Phacelia by both authors. Funding was provided in part by NSF GK12, NSF TREE, NSF GRF, and UC Berkeley Chancellor’s ee aye ee eee eee Phacelia subsect. Humiles Fellowships, and by research grants from the California Native Plant Society (CNPS Bristlecone Chapter, CNPS Marin County Chapter, and CNPS Orange County Chapter), Colorado Native Plant Society, Conservation Genetics Laboratory at SFSU, Lawrence R. Heckard Endowment Fund of the Jepson Herbar- ium, Nevada Native Plant Society, Southern California Botanists, SFSU Department of Biology, and UC Valentine Eastern Sierra Reserve to GKW. We thank Hartmut Hilger for directing our attention to the earlier tribe name on Jim Reveal’s INSG site. 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A new species of Solanum named for Jeanne Baret, an overlooked contributor to the history of botany. PhytoKeys 8:37-47. TORREY. 1871. Botany. Jn United States geological expolration [sic] of the Fortieth Parallel, S. Watson (ed.). Washington, DC. Voss, J. W. 1937a. A revision of the Phacelia Crenulata group for North America. Bulletin of the Torrey Botanical Club 64:81—96. . 1937b. A revision of the Phacelia Crenulata group for North America (Concluded). Bulletin of the Torrey Botanical Club 64:133—-144. WALDEN, G. K. 2010. Phylogeny of infrageneric relationships within Phacelia (Boraginaceae) in- ferred from chloroplast sequence data. M.S. Thesis, San Francisco State University, San Francisco, CA. AND R. PATTERSON. 2010. Howellanthus dalesianus, recognition of a new genus and species in tribe Phacelieae (Boraginaceae). Madrono 57:268—273. APPENDIX | CLASSIFICATION OF PHACELIA Phacelia Juss. Phacelia subg. Phacelia Phacelia sect. Phacelia Phacelia subsect. Phacelia Phacelia subsect. Humiles Walden & R. Patt. Phacelia sect. Baretiana Walden & R. Patt. Phacelia sect. Cosmantha (Nolte ex A. de Candolle) Benth. & Hook.f. Phacelia subsect. Cosmantha (Nolte ex A. de Candolle) Walden & R. Patt. Phacelia subsect. Bipinnatifidae (Small) Walden & R. Patt. Phacelia subsect. Cosmanthoides (A. Gray) Walden & R. Patt. Phacelia subsect. Dubiae (Small) Walden & R. Patt. Phacelia subsect. Ranunculacea Walden & R. Patt. N N MADRONO [Vol. 59 Phacelia sect. Eutoca (R. Br.) Benth. & Hook.f. Phacelia subsect. Eutoca (R. Br.) Walden & R. Patt. Phacelia subsect. Lineares (Rydb.) Walden & R. Patt. Phacelia subsect. Sericeae (Rydb.) Walden & R. Patt. Phacelia sect. Glandulosae (Rydberg) Walden & R. Patt. Phacelia sect. Gymnobytha (A. de Candolle) Benth. & Hook.f. Phacelia sect. Ramosissimae (Rydberg) Walden & R. Patt. Phacelia sect. Whitlavia (Harv.) Benth. & Hook.f. Phacelia subsect. Whitlaviae (Harv.) G. W. Gillett Phacelia subsect. Campanulariae G. W. Gillett Phacelia subg. Microgenetes (A. de Candolle) A. Gray Phacelia sect. Euglypta S. Watson Phacelia sect. Miltitzia (A. de Candolle) J. T. Howell Phacelia. sect. Pachyphyllae Walden & R. Patt. Phacelia subg. Pulchellae (Rydb.) Walden & R. Patt. MADRONO, Vol. 59, No. 4, pp. 223-229, 2012 DUDLEYA CRASSIFOLIA (CRASSULACEAE), A NEW SPECIES FROM NORTHERN BAJA CALIFORNIA, MEXICO MARK W. DODERO! AND MICHAEL G. SIMPSON Department of Biology, San Diego State University, San Diego, CA 92182 mdodero@reconenvironmental.com ABSTRACT Dudleya crassifolia is described as new. It belongs to a complex of taxa within subgenus Hasseanthus that have a white (occasionally pale yellow) corolla with a conspicuous, “‘musky-sweet”’ odor. It differs from related species of this complex in having a thicker petiole, a leaf blade that is only slightly wider than the petiole, and conspicuous dried leaf bases persisting on the caudex. This species is very rare, currently known only from a single population of scattered individuals (total area of coverage less than one hectare), on sandstone bluffs at Colonet Mesa of north coastal Baja California, Mexico. Key Words: Baja California, Crassulaceae, Dudleya, Dudleyva crassifolia, Hasseanthus, maritime chaparral, Mexico. The genus Dudleya Britton & Rose (1903), consists of approximately 33 species and 54 taxa (including infraspecies) of leaf-succulent perenni- als, native to California, Oregon, Arizona, Nevada, and Utah, USA, and to Baja California, Baja California Sur, and Sonora, Mexico (Wig- gins 1980; Kartesz 2011; McCabe 2012). Species of Dudleya differ from those of related genera, such as Sedum, in having axillary inflorescences (Moran 1942). Dudleya subgenus Hasseanthus, as traditionally treated (Moran 1950, 1951; Munz 1974; Bartel 1993; McCabe 2012) contains five species and six taxa: D. blochmaniae (Eastw.) Moran subsp. blochmaniae, D. blochmaniae (Eastw.) Moran subsp. insularis (Moran) Moran, D. brevifolia (Moran) Moran, D. multicaulis (Rose) Moran, D. nesiotica (Moran) Moran, and D. variegata (S. Watson) Moran. Species in subgenus Has- seanthus are characterized by their drought- deciduous leaf duration and hypogeous caudex, and range from San Luis Obispo Co., California to northern Baja California, Mexico. While attempting to relocate Baja California popula- tions of D. blochmaniae previously collected by the late Dr. Reid Moran, the first author discovered a distinct form of Dudleya at Colonet Mesa of north, coastal Baja California. We believe this form of Dudleya should be treated as a new species using a taxonomic (morphologic) species concept (Cronquist 1978, 1988), in which species are circumscribed based on the disconti- nuity of morphological features. ‘Present address: RECON Environmental, Inc., 1927 Sth Ave, San Diego, CA 92101. TAXONOMY Dudleya crassifolia Dodero & M. G. Simpson, sp. nov. (Figs. |, 2)—Type: MEXICO, Baja California, ca. 7 miles southwest of Colonet, reddish sandstone bluffs on mesa, maritime chaparral, associates: Adenostoma fascicula- tum, Agave shawii, Artemisia californica, Cea- nothus verrucosus, Cneoridium dumosum, Du- dleya attenuata subsp. a., Dudleya cf. ingens, Ephedra sp., Eriogonum fasciculatum, Ferocac- tus viridescens, Helianthemum scoparium, Rhus integrifolia, Salvia brandegeei, Stephanomeria sp., clay sub-soil, with upper layer of sand and surface iron concretions, 95 meters elevation, 31.00°N, 116.30°W, 3 June 1991, M. Dodero s.n. (holotype: SD; isotypes: BCMEX, RSA, iC). Diagnosis: Dudleya crassifolia is similar to D. blochmaniae subsp. blochmaniae, D. blochmaniae subsp. insularis, D. brevifolia, and D. nesiotica, but differs in having a thicker (2—3.3 mm) petiole, a leaf blade width at maturity only slightly greater than the petiole width (blade width:petiole width ratio 1.2—1.4), and conspicuously persistent dried leaf bases on the subterranean caudex. Description: Plants single to branched at base with up to 10 rosettes. Caudex cylindrical to irregular 1.0—5.5 cm long, 2—20 mm thick, with conspicuous, dried leaf bases persisting from multiple, previous seasons. Leaves in a rosette, 3-21, oblanceolate in mature plants to globose in juveniles, apex acuminate to acute to rounded (especially in juvenile plants), often mucronulate, 6-35 mm long, 3—11 mm wide, 1.5—7.5 mm thick, the petioles 0.9-12.4 mm long, 1.0—8.0 mm wide, 2—3.3 mm thick, the base truncate, 1-12 mm wide (narrower petioles are found in juvenile plants), leaves drought deciduous, bases persistent. Inflo- rescence 6—17 cm tall, ascending to inclined, |.5— N Nw & FIG. 1. MADRONO [Vol. 59 Habit shots, Colonet Mesa, Baja California, Mexico. A. Open, flat area of iron concretions, surrounded by mixed maritime chaparral. B—D. Dudleya crassifolia. B—C. Plants prior to flowering, showing one or more rosettes of thick, succulent leaves. D. Flower and buds of inflorescence. Note divergent, white corolla lobes. 3.5 mm thick at the base, bracts within 1—2 cm of the base. Lower bracts 7-19, ovate to widely ovate, acute, the lowermost 9-20 mm long, 6— 11 mm wide, 3—7 mm thick. Inflorescence axis usually with 2-3 ascending branches that re- branch 0-1 times, the ultimate cincinni 1-4 cm long, each with 2—10 flowers. Pedicels ascending, 1—3.5 mm long. Flowers with a characteristic musky odor. Calyx 3.5—6.5 mm wide, 3.5—6 mm long, lobes narrowly triangular to ovate, apex acute to acuminate, 2.5-4 mm long, 1.5—2.5 mm wide. Corolla 4-5 mm wide at the base, 14-17 mm wide at apex, basally connate 1-2 mm, lobes white, keel and base maroon, often suffused with maroon throughout, elliptic, 8-12 mm long, 3— 4.5 mm wide, ascending to divergent, apex acute. Stamen filaments 6-8 mm long, epipetalous, adnate 1-2 mm; anthers | mm long, 0.8 mm wide, red before dehiscence, yellow afterward. Nectar glands light yellow, 1.2 mm _ wide. Gynoecium 5—8 mm long, pistils erect to ascend- ing at anthesis, ovaries five, 4-6 mm long, styles 1-2 mm long. Fruits not described. Chromosome number: 7 = 34 (Dodero 1995). Distribution and Habitat: Dudleya crassifolia is endemic to the southern end of Colonet Mesa, southwest of Colonet, Baja California, Mexico (Fig. 3), in maritime chaparral, on reddish sandstone bluffs with a subsoil of clay covered with sand and iron concretions. Its total area of occurrence is less than one hectare. (See Discussion. ) Phenology: Dudleya crassifolia flowers from late May to June. Etymology: The specific eptithet, crassifolia, is from the Latin crassus “‘thick”’ and folius “leaf,” in reference to the thick, succulent leaves. Suggested common name: Thick Leaf Dudleya. Paratypes: MEXICO, Baja California, ca. 7 mi SW of Colonet, reddish sandstone bluffs on N N N 2012] DODERO AND SIMPSON: A NEW DUDLEYA SPECIES FROM MEXICO Fic. 2. Line drawings of Dudleya crassifolia from type population. A. Caudex, showing roots and apical rosette of leaves at apex. Note remains of old leaf bases below rosette. B. Inflorescence. Note succulent bracts along primary axis. C. Open flower. Note recurved corolla lobes, the upper half divergent. N N ON SN D. blochmaniae subsp. insularis D. blochmaniae subsp. b. oe California D. nesiotica ale D. brevifolia 100 km eA 3 1° ay v D. crassifolia © Fic. 3. Distribution map, from geo-referenced her- barium collections, of the taxa of the “blochmaniae” complex of Dudleya, subgenus Hasseanthus. Note wide distribution of D. blochmaniae subsp. blochmaniae (gray circles) and restricted distributions of D. blochmaniae subsp. insularis (white circles), D. brevifolia (white squares), D. nesiotica (white triangles), and the single known population of D. crassifolia (black star). mesa, maritime chaparral, associates: Adenos- toma fasciculatum, Agave shawii var. shawii, Artemisia californica, Ceanothus verrucosus, Cneoridium dumosum, Dudleya attenuata subsp. a., Dudleya cf. ingens, Ephedra sp., Eriogonum fasciculatum, Ferocactus viridescens, Helianthe- mum scoparium, Rhus integrifolia, Salvia brande- geei, Stephanomeria sp., clay sub-soil, with upper layer of sand and surface iron concretions, 95 m elev., 31.00°N, 116.30°W, 1 June 1992, M. Dodero s.n. (SBBG, SDSU, MEXU, UCR); towards the point of Punta Colonet, on the mesa, small sandstone outcrop, in maritime chaparral, associates: Adenostoma fasciculatum, Gambelia juncea [Galvezia j.], Cneoridium dumo- sum, Ceanothus verrucosus, Artemisia californica, Ferocactus viridescens, Agave shawii, Salvia bran- degeei, and Antirrhinum nuttallianum, ca. 84 m elev., 31.00°N, 116.31°W, 21 June 2008, Vander- plank C-66 (RSA, SD). DISCUSSION Morphological Distinctiveness Dudleya crassifolia is a distinctive entity of the genus that we feel warrants status as a separate species. It is a member of a group of species within subgenus Hasseanthus, this group also including D. blochmaniae subsp. blochmaniae, D. blochma- niae subsp. insularis, D. brevifolia, and D. nesiotica. These taxa, referred to here as the “blochmaniae complex,” share a white (rarely pale yellow in D. brevifolia and D. nesiotica) corolla MADRONO [Vol. 59 color and presence of a musky or “‘musky sweet” (McCabe 2012) flower odor, a combination that appears to be unique in the genus. All members of subgenus Hasseanthus are similar in juvenile leaf characteristics (Dodero 1995). In mature, adult leaves Dudleya crassifolia overlaps with members of the blochmaniae complex in blade width and petiole width, but in D. crassifolia the leaf blade is only slightly wider than the petiole (blade width:petiole width ratio 1.2—1.4), while in the other members of the complex the blade is two to more than four times as wide as the petiole (blade width:petiole width ratio 2.1—4.7) (Dodero 1995). Dudleya crassifolia also differs from the other four members of the blochmaniae complex in having significantly thicker petioles (2—-3.3 mm in D. crassifolia; ranging 0.4—-1.7 mm in the other four taxa; Dodero 1995). The other two members of subgenus Has- seanthus, Dudleya multicaulis and D. variegata, differ from D. crassifolia and the rest of the blochmaniae complex in having consistently yellow petals that lack the characteristic flower odor. Dudleya crassifolia also differs from D. multicaulis in its thicker, oblanceolate leaves with a smaller blade length to width ratio (2-3); in contrast, D. multicaulis has linear, more terete leaves with a much greater blade length to width ratio (6-11) (Dodero 1995). Dudleya crassifolia differs from D. variegata in the former’s greater petiole width and leaf thickness and smaller leaf- blade width to petiole width (1.2-1.4 in D. crassifolia versus ratio 2.6-4.1 in D. variegata). Finally, Dudleya crassifolia differs from all other species of subgenus Hasseanthus in having conspicuous, persistent dried leaves at the caudex base. We have to date seen no evidence for hybridization between D. crassifolia and other Dudleya species. According to a phylogenetic study using allozyme data (Dodero 1995), Dudleya crassifolia is most closely related to the D. blochmaniae subsp. blochmaniae populations that are found within a few miles of D. crassifolia (Fig. 3). These D. blochmaniae populations occupy patches of clay soils along with associates such as Chlor- ogalum parviflorum and Deinandra spp. Interest- ingly, the Dudleya blochmaniae populations in this general area of Baja California are known to be tetraploid (7 = 34) or hexaploid (n = 51) (Uhl and Moran 1953), the former identical to D. crassifolia (n = 34; Dodero 1995). In contrast, populations of D. blochmaniae in California are all diploid (n = 17) (Uhl and Moran 1953). Distribution, Evolution, Associates, and Geology Dudleya crassifolia is found approximately seven miles southwest of Colonet, Baja Califor- nia. It is apparently restricted to a few small, reddish sandstone bluffs of Colonet Mesa, 2012] defined as the area “‘south of the road to San Antonio del Mar” (Harper et al. 2011). This population consists of scattered individuals occu- pying an area estimated at less than one hectare. The bluffs where this species is found are covered with iron concretions. Just east of the sandy bluff is a partially stabilized dune formation with typical dune scrub (Holland and Keil 1995). An approximate distribution map of the taxa of the blochmaniae complex, assembled from geo- referenced data (BajaFlora 2012; CCH 2012; SEINet 2012) is seen in Fig. 3. Dudleya blochma- niae subsp. blochmaniae has a wide distribution, from coastal San Luis Obispo Co., California to coastal northern Baja California. The other taxa of Dudleya in this complex have a narrow distribution, D. blochmaniae subsp. insularis restricted to Santa Rosa Island (Santa Barbara Co.), D. brevifolia restricted to southwestern San Diego Co., D. nesiotica restricted to Santa Cruz Island (Santa Barbara Co,), and D. crassifolia restricted to Colonet Mesa of coastal northern Baja California (Fig. 3). A working hypothesis 1s that D. blochmaniae subsp. b. may represent a widespread, possibly paraphyletic species (after Olmstead 1995), from which the other taxa of the blochmaniae complex have diverged from isolated populations. This notion remains to be tested with molecular phylogenetic studies. Plant associates growing in the immediate vicinity of D. crassifolia include Adenostoma fasciculatum Hook & Arn., Agave shawii Engelm., Artemisia californica Less., Ceanothus verrucosus Nutt., Cneoridium dumosum (Torr. & A. Gray) Baill., Dudleya attenuata (S. Watson) Moran subsp. attenuata, Dudleya cf. ingens, Ephedra sp., Erio- gonum fasciculatum Benth., Ferocactus viridescens (Torr. & A. Gray) Britton & Rose, Helianthemum scoparium Nutt., Rhus integrifolia (Nutt.) W.H. Brewer & S. Watson, Salvia brandegeei Munz, and Stephanomeria sp. This community is best categorized as maritime chaparral, with surround- ing areas supporting Martirian succulent scrub, a more xerophytic vegetation type (Westman 1983). Dudleya crassifolia occurs in openings of this maritime chaparral. The associated plant species and red sandstone bluffs are apparently part of an ancient beach ridge formation similar to the sandstone bluffs found near Del Mar, San Diego Co., California, where D. brevifolia is found. Dudleya brevifolia occurs on bluff tops of the Linda Vista Forma- tion, which were formed as the sea receded during the Pleistocene. The surface of these bluffs is covered with small iron concretions similar to the one at Colonet Mesa. Herbivory, Pollination, and Seed Dispersal Observations indicate that herbivory, presum- ably by rabbits (rabbit scat is frequently seen on DODERO AND SIMPSON: A NEW DUDLEYA SPECIES FROM MEXICO 227 the bluff) and small rodents, appears to be significant. In November 1992, before any rain had fallen and conditions were very dry, many dormant plants had been excavated by animals and eaten. The leaves of actively growing plants were eaten by animals during the winter growing season. In cultivation, plants in which the top of the tuberous caudices were exposed above the soil surface, and which appeared to be dead, are capable of re-sprouting from beneath the soil surface, often forming multiple rosettes. Potential pollinators were observed and captured. Two species of wasp, Stenolia dupli- cata and Bembix occidentalis, both members of the sand wasp group (family Crabronidae, subfamily Bembicinae), were seen visiting flow- ers. Typically these wasps feed on flies, but they will supplement their diet with pollen (Borror and White 1970; Borror et al. 1989). Another potential pollinator, the soft-winged flower bee- tle, Dasytes sp. (family Melyridae, subfamily Dasytinae) lives in the flowers. These beetles are usually found at the base of the carpels near the nectar glands. Seed dispersal appears to be accomplished mostly by wind and water movement. Seeds sprinkled on the soil surface during a 15—20 knot wind were observed to move along with grains of sand. If moisture and temperature conditions are optimal, germination of D. crassifolia seeds 1s rapid, often occurring in less than 72 hours. Recruitment of seedlings appears to be highest in areas of moderate soil deposition and underneath shrubs where seeds are transported by the wind (Dodero, personal observations). Conservation The Colonet region is estimated to have the highest plant diversity in Baja California (Harper et al. 2011). The region has 435 documented vascular plant taxa, of which 383 are native to Baja California, and 52 are endemic or nearly endemic to the state or peninsula (Harper et al. 2011). Eighteen of these taxa are listed by the California Native Plant Society as 1B, “rare, threatened, or endangered in California and elsewhere” (CNPS 2012). Three of these taxa are on the Mexican NOM 059 list of protected taxa; five of these taxa are narrow endemics, known only from the Colonet region (Harper et al. 2011). Of the five taxa that are local endemics to the Colonet region, Dudleya crassifolia is perhaps the rarest and most geographically restricted, being found at a single locality on sandstone bluffs at Colonet Mesa. Its rarity and limited distribution watrant focused protective measures. Unfortu- nately, Colonet Mesa and the surrounding region is threatened by a pending proposal from the Mexican government to construct a seaport 228 (which would be accompanied by massive infra- structure and a large, projected growth in human population) just below the southern tip of the mesa at Punta Colonet. This project could MADRONO [Vol. 59 seriously impact this unique and diverse region without major conservation efforts to protect sensitive habitat (Clark et al. 2008; Harper et al. 2011). Taxonomic Key A revised key to species of subgenus Hasseanthus, modifed from McCabe (2012), 1s presented below. 1. St gen below surface, not elongate, gen simple, unbranched; leaves deciduous, vernal, generally petioled (barely in D. crassifolia) (subg. Hasseanthus) 2. Flower odor absent; corolla consistently yellow 3. Leaves 4-15 cm long, linear, + narrowed above base, tip sharply acute; petals basally connate ee A ee ee ee ee ee Me Gh es ARE Sek taetrecaniy At on Mane an ty ae, Ree D. multicaulis 3’ Leaves 1—7 cm long, oblanceolate to spoon-shaped, strongly narrowed above base (to gen 0.5— 3 mm wide), tip acute to obtuse; petals basally connate 0.5-lmm................ D. variegata 2’ Flower odor musky-sweet; corolla white to white-maroon (rarely pale yellow in D. brevifolia and D. nesiotica) 4. Leaf blade only slightly wider than petiole (ratio 1.2—1.4), petioles 2—3.3 mm thick; caudex with conspicuous, dried leaf bases persisting from multiple, previous seasons........... D. crassifolia 4’ Leaf blade two to more than four times as wide as the petiole (ratio 2.1-4.7), petiole 0.4—1.7 mm thick; caudex lacking conspicuous, dried leaf bases from previous seasons 5. Petals ascending, 7-14 mm, 3.5—5.5 mm wide, fused 1—2 mm; fr ascending; If base 3-12 mm WIdGs 2 acl os oe ee ae ee D. nesiotica 5’ Petals spreading, 5-10 mm, 2-4 mm wide, fused gen <1 mm; fr spreading; If base 14 mm wide 6. Lower bracts <1.5x longer than wide; lvs 7-15 mm, + spheric to spoon-shaped; petiole WALEOW 4c 4 seas ee ee 6’ Lower bracts >2x longer than wide; lvs 10— 60 mm, + petiole = marrow. «2... 2... 24-.. 7. Lvs gen <12, not to + glaucous 7’ Lvs gen >15, glaucous or + so Digs hase Bk aus Aah @ Slate wees Pau Sen Sal D. brevifolia oblanceolate to club-shaped; Oe ee ee ee ee re ee Ne ee D. blochmaniae Te ee ee subsp. blochmaniae ee ee eee ee Le ee ree subsp. insularis 1’ St gen above surface, often elongate, often branched; lvs gen evergreen, + not petioled (remaining Dudleya species) ACKNOWLEDGMENTS We thank Jim Berrian and Robyn Garcia for providing transportation and assisting in field work, Bruce and Karen April for help in the field, Robyn Garcia for providing the illustrations, and Kim Marsden for assistance in field work and moral support. Dave Truesdale was indispensable in the allozyme work cited here. Finally, we thank three anonymous reviewers for valuable comments made in review of the manuscript. LITERATURE CITED BAJAFLORA. 2012. The flora of Baja California. Specimen data obtained from a combined multi- herbarium specimen database called the Baja California Botanical Consortium hosted by the San Diego Natural History Museum. San Diego, CA.Website: http://www. bajaflora.org [accessed 18 April 2012]. BARTEL, J. A. 1993. Dudleya. Pp. 525-530 in J. C. Hickman (ed.), The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. BORROR, D. J. AND R. E. WHITE. 1970. A field guide to the insects of America north of Mexico. Houghton Mifflin, Boston, MA. ———, C. A. TRIPLEHORN, AND N. F. JOHNSON. 1989. An introduction to the study of insects, 6th ed. Saunders College Publishing, Philadelphia, PA. BRITTON, N. L. AND J. N. ROSE. 1903. New or noteworthy North American Crassulaceae: Du- dleya Britton & Rose, gen. nov. Bulletin of the New York Botanical Garden 3:12—28. CONSORTIUM OF CALIFORNIA HERBARIA (CCH). 2012. Website: http://ucjeps.berkeley.edu/consortium [ac- cessed 18 April 2012]. CLARK, K. B., M. DODERO, A. CHAVEZ, AND J. SNAPP- Cook. 2008. The threatened biological riches of Baja California’s Colonet Mesa. Fremontia 36:3—10. CALIFORNIA NATIVE PLANT SOCIETY (CNPS). 2012. Inventory of rare and endangered plants (online edition, v8-Ola). California Native Plant Society, Sacramento, CA. Website: http://www.rareplants. cnps.org/ [accessed 23 February 2012}. CRONQUIST, A. 1978. Once again, what is a species? Pp. 3— 20 in J. A. Ramberger (ed.), Biosystematics in agriculture. Allanheld & Osmun, Montclair, NJ. 1988. The evolution and classification of flowering plants, 2nd ed. New York Botanic Garden, New York, NY. DoDERO, M. W. 1995. Phylogenetic analysis of Dudleya subgenus Hasseanthus (Crassulaceae) using mor- phological and allozyme data. M.S. thesis, San Diego State University, San Diego, CA. HARPER, A. B., S. VANDERPLANK, M. DODERO, S. MATA, AND J. OCHOA. 2011. Plants of the Colonet region, Baja California, Mexico, and a vegetation map of Colonet Mesa. Aliso 29:25—-42. HOLLAND, V. L. AND D. J. KEIL. 1995. California vegetation. Kendall/Hunt, Dubuque, IA. KARTESZ, J. T. 2011. The biota of North America program (BONAP). North American Plant Atlas. Chapel Hill, NC. Website http://www.bonap.org/). [maps generated from Kartesz, J. T. 2010. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP)], [accessed 01 December 2011]. | \ 2012] McCABE, S. W. 2012. Dudleya. Pp. 666-673 in B. G. Baldwin, D. H. Goldman, D. J. Keil, R. Patterson, T. J. Rosatti, and D. H. Wilken, (eds.), The Jepson manual: vascular plants of California, 2nd ed. University of California Press, Berkeley, CA. MORAN, R. 1942. The status of Dudleya and Stylo- phyllum. Desert Plant Life 14:149—157. . 1950. Notes on Hasseanthus 1. Desert Plant Life 22:6—-82. . 1951. Notes on Hasseanthus I. Desert Plant Life 22:101—105. Munz, P. A. 1974. A flora of Southern California. University of California Press, Berkeley, CA. DODERO AND SIMPSON: A NEW DUDLEYA SPECIES FROM MEXICO 229 OLMSTEAD, R. G. 1995. Species concepts and plesio- morphic species. Systematic Botany 20:623—630. SOUTHWEST ENVIRONMENTAL INFORMATION NETWORK (SEINET). 2012. Website: http://swbiodiversity.org/ seinet/index.php [accessed 18 April 2012]. UHL, C. H. AND R. MORAN. 1953. The cytotaxonomy of Dudlevya and Hasseanthus. American Journal of Botany 40:492—502. WESTMAN, W. E. 1983. Xeric Mediterranean-type shrub- land associations of Alta and Baja California and the community/continuum debate. Vegetatio 52:3—19. WIGGINS, I. L. 1980. Flora of Baja California. Stanford University Press, Stanford, CA. MADRONO, Vol. 59, No. 4, p. 230, 2012 NOTEWORTHY COLLECTION CALIFORNIA CHIMAPHILA MENZIESII (R. Br. ex D. Don) Sprengel (ERICACEAE). —Humboldt Co., Lanphere Dunes Unit, Humboldt Bay National Wildlife Refuge, 5 m, Tyee City quadrangle Section 23, SENW, 10S 0403652 E., 4526990 N., one population of 50 plants was found growing ina 3 X 3 m area ina coniferous dune forest, in unvegetated duff under Morella californica with Pinus contorta subsp. contorta, Picea sitchensis, Goodyera oblongifolia, Vaccinium ovatum, Frangula purshiana, and the terricolous moss Kindbergia oregana. Plants were less than 10 cm. tall, 21 February 2011, Adam N. Canter 003 (HSC). One coastal population previously noted (03 June 1979, Seacat and Seymour 193 (HSC)) was relocated; Mendocino Co. quadrangle Section 28 SWSE, 10S 0434180 4350344, 5 m, approximately 1.5 mile upriver, south side, in a mixed conifer forest, Pseudotsuga menziesii dominant stand with adjacent Tsuga hetero- phylla, Abies grandis, and Sequoia sempervirens. Under- story consisted of Lithocarpus densiflorus, Rhododendron macrophyllum, Vaccinium parvifolium, and Polystichum munitum. Herbaceous layer included Oxalis oregana, Disporum smithii, Goodyera oblongifolia, Vancouveria spp., Berberis aquifolium, Hierochloe occidentalis, and Kindbergia oregana. Population +/— 50 individuals, 10 x 10 m area, 27 November 2011, Adam N. Canter, Ana L. R. Canter 001 (HSC). A population of 18 individuals was found approxi- mately 1 km south of the initial detection discussed above, (Adam N. Canter 003); Section 23 SESW, 10S 0403713 E., 4526777 N., in a more Picea sitchensis dominated patch, within the same plant community, 21 February 2011. No physical collection was made due to the conservation concerns of this small population. A third population of 7 individuals was detected further south at the Ma-le’] Dunes Unit, under dense Vaccinium ovatum with Chimaphila umbellata and Listera cordata as community associates in addition to those at the other sites, Section 26 NESW, 10S 0403474 E., 4526046 N., 01 December 2011, Adam N. Canter 002 (HSC). Previous knowledge. Chimaphila menziesii is treated by Wallace and Haber (2012) as uncommon, occurring in montane conifer forest, 1000—2500 m, and flowering June-August. The lower elevational range is closer to 400-500 m, according to the most recent list of records for this species in California provided by the California Consortium of Herbaria (2012). Coastal populations have been noted in Haida Gwaii (Queen Charlotte Islands), British Columbia, first detected in 1999 (Smith and Buttler 1999; Jan Oord, Laskeek Bay Conservation Society, personal communication) and one isolated population on the Big River Estuary, Mendocino Co. California (03 June 1979, Seacat and Seymour 193 (HSC)). Significance. This is the second report of Chimaphila menziesii in the hypermaritime coastal zone, south of British Columbia, and the second documented detec- tion in the North Coast subregion of Northwestern California. Plants were not observed flowering until October, and continued to flower through December 2011. —ADAM N. CANTER, U.S. Fish and Wildlife Service, Humboldt Bay National Wildlife Refuge, 6800 Lanphere Rd., Arcata, CA 95521. acanter@vt.edu. LITERATURE CITED CALIFORNIA CONSORTIUM OF HERBARIA. Website: http://ucjeps.berkeley.edu/consortium/ [accessed 20 March 2012]. SMITH, J. AND I. BUTTLER. 1999. First record of Menzies’ pipsissewa, Chimaphila menziesii, on Haida Gwaii. Pp. 29 in A. J. Gaston (ed.), Laskeek Bay Research 9. Laskeek Bay Conservation Soci- ety, Queen Charlotte City, B.C. WALLACE, G. D. AND E. HABER. 2012. Chimaphila in Jepson flora project (v. 1.0), Jepson eFlora, Regent of the University of California, Berkeley, CA.Web- site: http://ucjeps.berkeley.edu/IJM.html [accessed 02 February 2012]. MADRONO, Vol. 59, No. 4, p. 231, 2012 NOTEWORTHY COLLECTION CALIFORNIA CAREX NARDINA Fries (CYPERACEAE ).—Siskiyou Co., Klamath National Forest, Marble Mountain Wilderness, slopes of Black Marble Mountain, 41.57836°, —123.20355° (NAD 83), 2110 m/6905 ft., 13 August 2011, York 3059 (HSC, OSU). A rare, clumped perennial with a single spikelet growing on limestone shelves on a large, north-facing outcrop with Achillea millefolium, Angelica arguta, Arnica sp., Campanula rotundifolia, Cystopteris fragilis, Oxyria digyna, Polystichum lonchitis, and Saxifragiopsis fragar- ioides. Siskiyou Co., Klamath National Forest, Marble Mountain Wilderness, slopes of Kings Castle, approx. 5 km north of other known population (York 3059) for, CA, 4161475", —123.22112° (NAD 83); 2175 m/7150 ft., 8 September 2011, York 3087 (HSC, JEPS, OSU). A rare, clumped perennial with a single spikelet growing on limestone shelves on a large, northeast- facing outcrop with Achillea millefolium, Angelica arguta, Arnica mollis, Campanula rotundifolia, Cystop- teris fragilis, Pellaea breweri, Polemonium pulcherri- mum, and Polystichum lonchitis. Previous knowledge. Carex nardina is a circumpolar species occurring throughout the northern portion of North America into Eurasia. According to Murray (2002), there has been much written about the variation in C. nardina with little resolution. Some authors have considered some or all the North American material to be C. n. subsp. hepburnii (Boott) A. Love, D. Love & B. M. Kapoor or C. hepburnii Boott, but no clear geographic limits occur. Wilson et al. (2008) recognize C. nardina as occurring mostly in the northeastern portion of Oregon. The single known southern Oregon population, representing York 288/ (OSU) from Mount Thielsen, 8 August 2005, is the closest known occur- rence to the Marble Mountain Wilderness collection. It is nearly 190 air-kilometers away. There are no known populations of C. nardina in the bordering states of Nevada and Arizona. Significance. This is the first documented collection from California. —DANA A. YORK, Senior Environmental Planner, California Department of Transportation, 1656 Union Street, Eureka, CA 95501. Dana_York(@dot.ca.gov. LITERATURE CITED MURRAY, D. F. 2002. Carex nardina. Pp. 567-9 in Flora of North America Editorial Committee (eds.), Flora of North America north of Mexico, Volume 23, Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford University Press, New York, NY. WILSON, B. L., R. BRAINERD, D. LYTJEN, B. NEw- HOUSE, AND N. OTTING. 2008. Field guide to the sedges of the Pacific Northwest. Oregon State University Press, Corvallis, OR. MADRONO, Vol. 59, No. 4, pp. 232-233, 2012 NOTEWORTHY COLLECTIONS CALIFORNIA NAJAS MINOR Allioni (HY DROCHARITACEAE).— Alameda Co., Lake Del Valle Park, southeast end of Lake Del Valle, along east side of peninsula extending northward from southeast side of West Swim Beach swimming area, 37°34'47.7"N, 121°41'48.5”W, drifting fragments fairly common along the margins of the lake in shallow water, flowers and some fruits present, 23 July 2011, Donald H. Les 1029 [with Hamid Raczifard] (CONN). This California material was collected under the East Bay Regional Parks permit #596 held by the University and Jepson Herbaria, University of Califor- nia, Berkeley. Alameda Co., San Francisco Bay Area Pond in Del Valle Park, 7000 Del Valle Rd, Livermore, 07 August 2003, K. Simmons PDR 1283308 (CDA) [misidentified as Najas marina L]. Alameda Co., San Francisco Bay Area Pond in Del Valle Park, 7000 Del Valle Rd, Livermore, 07 August 2003, K. Simmons PDR 1283309 (CDA) [misidentified as Najas marina L]. Previous knowledge. Najas minor was unknown in California prior to our collection and a search of the University of California-Berkeley herbarium failed to disclose any material. However, a survey of herbarium material from CDA disclosed three specimens identified as Najas marina, which were collected from the same locality (Lake Del Valle, CA) in 2001 and 2003. We confirmed the identity of the specimen collected in 2001 (J. R. Willson PDR P199008-C, [CDA]) as Najas marina; however, the two specimens collected in 2003 were misidentified and actually represent specimens of Najas minor. Consequently, the 2003 records provide the earliest verification of this nonindigenous species in the state of California. The closest known locality of N. minor occurs in Texas, approximately 2400 km south- eastward of the California occurrence (see Texas Noteworthy Collection). An early paper dealing with dermatitis in California rice field workers suggested that the malady might be caused by contact with the leaf “thorns” of the water plant Najas minor (Alderson and Rawlins 1925). However, the authors clarified that Najas minor did not occur in California, and that this ailment had been reported in the European literature. Certainly it is highly unlikely that this species would have been known from California a decade before the first verified occurrence in North America (Wentz and Stuckey 1971) and preceding by 78 years the first known voucher specimen from the state. However, Najas species (e.g., N. graminea) are known to occur as rice field weeds in both Australia and California as a result of contaminated planting stocks (McIntyre and Barrett 1985) and it is intriguing to think that the plants could have escaped detection for so long in such a thoroughly botanized region. Significance. Najas minor is a nonindigenous, invasive aquatic plant introduced to North America from Eurasia at some point before 1932 (Wentz and Stuckey 1971). Because of its nuisance status and expanding distribution, this record raises concern for the possible proliferation of new sites for this weedy plant in the western United States, where it fortunately has not yet established. The source of introduction for N. minor in California is unknown. However, because this species is known to spread from rice fields, it should be searched for thoroughly not only in the Sacramento Valley, but also throughout other rice-growing portions of the State. —DONALD LES, Department of Ecology & Evolu- tionary Biology, University of Connecticut, 75 North Eagleville Road, Storrs, CT 06269-3043; NICHOLAS TIPPERY, Department of Biological Sciences, University of Wisconsin-Whitewater, Whitewater, WI 53190-1790; HAMID RAZIFARD, Department of Ecology & Evolu- tionary Biology, University of Connecticut, 75 North Eagleville Road, Storrs, CT 06269-3043. les@uconn.edu. LITERATURE CITED ALDERSON, H. E. AND A. G. RAWLINS. 1925. Rice workers’ dermatitis. California and Western Med- icine 23:42-45, MCINTYRE, S. AND S. C. H. BARRETT. 1985. A comparison of weed communities of rice in Australia and California. Proceedings of the Ecological Society of Australia 14:237—250. WENTZ, W. A. AND R. L. STUCKEY. 1971. The changing distribution of the genus Najas (Najada- ceae) in Ohio. Ohio Journal of Science 71:292—302. TEXAS NAJAS MINOR Allioni (HY DROCHARITACEAE).— Bastrop Co., Lake Bastrop Park, boat launch site at the northwest end of the lake, 30°09'55.3”N, 097°16'49.0’"W, abundant along the pier in 1—2 m of water, with Najas guadalupensis and Hydrilla verticillata, 11 July 2010, Donald H. Les 843 and Nicholas P. Tippery 320 (CONN). Previous knowledge. This material represents the first verified record of N. minor in Texas. The Texas Parks and Wildlife Department (TPWD 2012) lists only “marine naiad’’ and “‘southern naiad’” among the aquatic vegetation of Lake Bastrop. We confirmed the existence of southern naiad (N. guadalupensis) at the site; however, we found no evidence of marine naiad (Najas marina) and suspect that N. minor had been misidentified as the latter because of their similarly toothed leaves. The strongly toothed leaves of WN. marina differ from all other North American species except N. minor, and this character often is used as a distinguishing trait for the former in keys written for areas where N. minor is not known to occur. Because of | its proximity in Louisiana and Oklahoma, Diggs et al. (2006; p. 676) concluded that Najas minor **... would thus not be unexpected in East TX.”’ Our collection substantiates that prediction. Significance. This record confirms the continued westward spread of this invasive aquatic plant in North America and provides the first verified floristic record for the species in the state of Texas. This locality also represents the most westward extension of N. minor in North America with the exception of the highly disjunct material from California, which is described above. A | Nw WwW ies) 2012] NOTEWORTHY COLLECTIONS re-examination of “‘Najas marina’ specimens from LITERATURE CITED Texas is suggested, given that it is likely to disclose : additional, earlier records of N. minor from the state. DicGs, Jr., G. M., M. D. REED, R. J. O"KENNON, AND B. L. Lipscoms. 2006. Illustrated flora of East —DONALD Les, Department of Ecology & Evolu- Texas. Austin College Center for Environmental tionary Biology, University of Connecticut, 75 North Studies and the Botanical Research Institute of Eagleville Road, Storrs, CT 06269-3043; NICHOLAS Texas, Austin, TX. TIPPERY, Department of Biological Sciences, University TEXAS PARKS AND WILDLIFE DEPARTMENT (TPWD). of Wisconsin-Whitewater, Whitewater, WI 53190-1790: 2012, Lake Bastrop. Texas Parks and Wildlife HAMID RAZIFARD, Department of Ecology & Evolu- Department, Austin, TX. Website http://www. tionary Biology, University of Connecticut, 75 North tpwd.state.tx.us/fishboat/fish/recreational/lakes/ Eagleville Road, Storrs, CT 06269-3043. les@uconn.edu. bastrop/ [accessed 15 June 2012]. MADRONO, Vol. 59, No. 4, p. 234, 2012 PRESIDENT’S REPORT FOR VOLUME 59 Dear CBS member, This year, 2012, has been a transitional year for the Society. We passed our 99th birthday as a Society on April 13, 2012. To celebrate our Centennial in April 2013, we are organizing a symposium in Berkeley, California, with invited speakers and a special banquet, in conjunction with the 23rd graduate student meetings. We have also planned, and have conducted, a series of field trips in honor of the 100th birthday. Please see www.calbotsoc.org for all the information on the Centennial. This year we signed an agreement with JSTOR (an online archiving system) to place all of our Madrofio back issues online. Current issues are already online through BioOne.2. We are moving toward having the current online issues easily available to members, so look forward to that. One of the journal’s editors, Richard Whitkus, has announced that he is stepping down after several critical years. We all are thankful to have had Rich during the time the journal converted to an online manuscript submission and tracking process and rapidly caught back up on schedule. For 2012, we did not hold a separate banquet meeting, but instead sponsored the banquet speaker for the California Native Plant Society’s January Conservation Conference in San Diego. Peter Raven, Director Emeritus of the Missouri Botanical Garden, was our invited speaker, and he gave a wonderful talk reflecting on the history of Western North American botany and what the future may hold. The Council has worked particularly hard for you this year and I would especially like to thank Andrew Doran, Dean Kelch, Kim Kersh, Staci Markos, Anna Larsen, Matt Ritter, Tom Schweich, Ellen Simms, Michael Vasey, and Rich Whitkus. All of our Council members have been critically important this year. We also welcome Genevieve Walden from UC Berkeley as the new graduate student representative; Genevieve will be organizing the 2013 graduate student meetings. Reaching 100 years as a Society has moved the Council into a reconsideration of the role of the California Botanical Society. The founding goals of supporting botanical research and communicating that broadly have been largely achieved, but California and western North America are no longer characterized by low populations and difficult travel. Journeying across our mountain ranges may take some time, but we no longer consider them expeditions. Electronic communication has rapidly changed the way scientists communicate and we are transitioning as fast as we can afford. Because our membership base is the foundation of the Society, we would like to hear from you about what roles you see for the Society in the future. Please contact any of the Council members and let us hear from you. Of course, increasing our membership is always a priority, so please continue to encourage your colleagues to join us and to publish in Madrono. This is especially true of our younger colleagues. Moving online should make us more attractive to the younger cohorts of botanists more accustomed to this format. Lastly, I regret to note the passing of Dr. John O. Sawyer (1939-2012) for whom our 2008 volume of Madrono was dedicated. John was a stellar plant ecologist who joined the faculty at Humboldt State University in 1966 and remained there until he retired. He mentored many graduate students who are now professional botanists, was a president of the CNPS and founder of its North Coast Chapter, and was instrumental in preparing two editions of the Manual of California Vegetation. His humor, deep passion for and knowledge of the vegetation of northwestern California will be sorely missed. V. Thomas Parker December 2012 MADRONO, Vol. 59, No. 4, p. 235, 2012 EDITORS’ REPORT FOR VOLUME 59 We are pleased to report the publication of Volume 59 of Madrono by the California Botanical Society (CBS) in 2012. This year we were successful in bringing the publication of Madrono back on schedule and we appreciate the efforts of our reviewers and contributors in this effort. The average turn-around time between initial submission and publication has been reduced to less than eight months and we hope that Madrono continues to be viewed as the best outlet for western botanists to publish their work in a timely fashion, and reach an interested and relevant audience. The Madrono page on the CBS web site has instructions for online submission of new manuscripts, digital versions of recent issues through BioOne, and the eighty-year index. The efforts of numerous individuals are critical to the continued quality and timeliness of the journal. Among these, our Noteworthy Collections editor, Dieter Wilken; Steve Timbrook who has long provided the volume Index and Table of Contents; Annielaurie Seifert at Allen Press; and the enthusiastic support of the CBS executive council. Finally, we are extremely grateful to our contributors for their interesting and insightful manuscripts, and our reviewers who take time from their busy schedules to assess the quality of submitted work. This year we received 31 new manuscripts and 26 were accepted for publication. Several manuscripts were also carried over from the previous year. The 59th volume includes 11 Articles (including Notes), six New Species, four Noteworthy Collections, two Book Reviews, and two Points of View. We are also excited to present in this volume the special issue Flora of the Carquinez Strait Region, Contra Costa and Solano Counties, California by Dean G. Kelch and Andrew Murdock. This beautifully done flora will be useful for western botanists in the San Francisco Bay Area and beyond. We appreciate the submission of a variety of different manuscripts across the spectrum of botanical sciences and anticipate continued submissions of novel and exciting work. We also hope to see the Points of View become a regular feature presenting insightful discussion of issues pertinent to botanists, managers, and policy makers. The coming year will be a once 1n a century event as the CBS celebrates our centennial year. The CBS is hosting the Centennial Symposium “Botanical Frontiers: Past and Future’’, at the University of California, Berkeley in April 2013. Please visit the CBS website (calbotsoc.org) for details about upcoming centennial celebration events including field trips to Mount Tamalpais and Mount Diablo, a symposium, banquet, and graduate student meeting. We are hoping that issues 3 and 4 of the upcoming volume will feature papers generated from presentation given at the Centennial Symposium. We also look forward to the second issue of the 60th volume—a special issue on tanoak (Notholithocarpus densiflorus). This issue will feature papers by a community of researchers studying the distribution, genetics, utilization and natural history of tanoak. The issue will be a synthesis of work presented during a session at the 5th Sudden Oak Death Science Symposium in June 2012 called ““What are we trying to save? Tanoak history, values and ecology”’. As Editors, we have enjoyed our interactions with contributors and reviewers this past year, and we look forward to an exciting, productive, and momentous centennial year for western botany. Matt Ritter Richard Whitkus December 2012 MADRONO, Vol. 59, No. 4, p. 236, 2012 REVIEWERS OF MADRONO MANUSCRIPTS 2012 Paul Beardsley Elizabeth Braker Leo Bruederle James Cohen Tom Daniel Naomi Fraga Timothy Griffith Lisa Grubisha Richard Halse Dylan Hannon Chris Havran Christina Hazard Kathleen Kay Jon Keely Ron Kelley Byron Lamont Thomas Marcussen Bruce Maslin Stephen McCabe Nicole Molinari Nancy Morin Tom Parker Robert Patterson Jon Rebman Michael Simpson Shannon Still Dean Taylor Mike Vasey Michael Vincent Mark Waldrop Gary Wallace Dieter Wilken Nina Wurzburger Jenn Yost MADRONO, Vol. 59, No. 4, pp. 237-238, 2012 INDEX TO VOLUME 59 Classified entries: major subjects, key words, and results; botanical names (new names are in boldface); geographical areas; reviews, commentaries. Incidental references to taxa (including most lists and tables) are not indexed separately. Species appearing in Noteworthy Collections are indexed under name, family, and state or country. Authors and titles are listed alphabetically by author in the Table of Contents to the volume. Alpine plants (see Yellowstone Nat. Park) Annual plants, effects of recurrent fire, 14. Boraginaceae (see Eriodictyon and Phacelia) Botanical code changes, implications for western No. Am. botany, 169. Bryaceae (see Ptychostomum) California: Carex albida and C. lemmonii, relationship, 171; effects of recurrent fire on annual plants of Colorado Desert, 14; Eriogonum evanidum, rediscovery, 150; flora of Carquinez Strait, 47; Grassland vegeta- tion, response to removal of sheep on Santa Cruz Id., 190; Vaccinium parvifolium, genetic structure, 196; Viola pedunculata, genetic structure and outcrossing rates, 181. New taxa: Calystegia binghamiae, 25; Eriodictyon traskiae var. smithii and E. t. var. traskiae, 28; Ptychostomum pacificum, 156; Rhododendron occi- dentale var. californicum, 140. Noteworthy collection: Carex nardina, 231; Chimaphila menziesii, 230; Najas minor, 232; Trifoloium tricho- calyx, 167. Calystegia: C. sepium subsp. binghamiae, rediscovery and status, 25. New taxon: C. binghamiae, 25. Carex albida and C. lemmonii, relationship, 171; C. nardina, noteworthy collection from CA, 231. Carquinez Strait, CA, flora, 47. Ceanothus cuneatus var. cuneatus and C. roderickii on gabbro soils, 1. Chapparal, postfire regeneration, 109. Chasmogamy (see Viola) Chimaphila menziesii, noteworthy collection from CA, 230. Chromosome counts (see Mimulus) Cleistogamy (see Viola) Convolvulaceae (see Calystegia) Crassulaceae (see Dudleya) Cyperaceae (see Carex) Dudleya crassifolia, new sp. from No. Baja Calif., MEXICO, 223. Editors’ Report for Vol. 59, 235. Ericaceae: Rhododendron: R. occidentale, morphological and isoenzyme variation, 128; Vaccinium parvifolium, genetic structure, 196. Noteworthy collection of Chimaphila menziesii from CA, 230. Eriodictyon traskiae var. smithii and E. t. var, traskiae, new taxa, 28. Eriogonum evanidum, rediscovery, 150. Fabaceae (see Trifolium) Feral sheep (see Grassland vegetation) Fire: Postfire chaparral regeneration under mediterra- nean and non-mediterranean climates, 109; recurrent fire, effects on annual plants of Colorado Desert, CA, 14. Floras: Carquinez Strait region, Contra Costa and Solano cos., CA, 47; alpine Mt. Washburn, Yellow- stone Nat. Park, WY, 2. Gabbro soils, |. Grassland vegetation, response to removal of sheep on Santa Cruz Id., CA, 190. Hydrocharitaceae (see Najas) Hydrophyllaceae (see Eriodictyon and Phacelia) Keys: CA members of Carex sect. Aulocystis, 177; Dudleya subg. Hasseanthus, 228; Phacelia subdivisions, 218; Rhododendron occidentale varieties, 140. Linanthus pungens subsp. hazeliae, new comb. from ID and OR, 163. Meadows, montane: Spatial autocorrelation of vege- tation in Sierra Nevada, CA and western NV, 143. MEXICO (see Dudleya) Mimulus: M. sookensis, new allotetraploid sp. derived from M. guttatus and M. nasutus, 29. Noteworthy collection: M. 166. Najas minor, noteworthy collections from CA and TX, 2am. clivicola from MT, Phacelia: Nomenclatural subdivisions new combinations and statuses, 211. Phrymaceae (see Mimulus) Polemoniaceae, (see Linanthus) Polygonaceae (see Eriogonum) Polyploidy (see Mimulus) President’s Report for Vol. 59, 234. Ptychostomum pacificum, new fen sp., 156. including Reviews: Northwest California: A Natural History by John O. Sawyer, 44; Research & Discovery in Vernal Pool Landscapes by D.G. Alexander and R. A. Schlising, 164. Rhamnaceae (see Ceanothus) Rhododendron: R. occidentale, morphological and isoen- zyme variation, 128. New taxon: R. occidentale var. californicum, 140. Santa Cruz Id, CA (see Grassland vegetation) Sierra Nevada, CA and western NV (see Meadows, montane) Spatial autocorrelation (see Meadows, montane) 238 MADRONO [Vol. 59 Texas (see Najas) Violaceae (see Viola) Trifoloium trichocalyx, noteworthy collection from CA, 167. Wilken, Dieter H., dedication of Vol. 59 to, 239 Wyoming (see Yellowstone Nat. Park) Vaccinium parvifolium, genetic structure, 196. Viola pedunculata, genetic structure and outcrossing rates, Yellowstone Nat. Park, Mt. Washburn vascular alpine 181. flora, 2. MADRONO, Vol. 59, No. 4, pp. 239-240, 2012 DEDICATION DIETER H. WILKEN The California Botanical Society dedicates Volume 59 to Dr. Dieter H. Wilken, Director of Conservation at the Santa Barbara Botanic Garden. Born in 1944, Dieter grew up in Los Angeles. He attended California State University, Los Angeles, where he received his Bachelor’s degree in 1967. He then moved to a Ph.D. program at the University of California, Santa Barbara, where he worked under the direction of Dale Smith. Dieter earned his Ph.D. in 1971, and his dissertation was entitled A biosystematic study of the genus Hulsea (Asteraceae: Helenieae ). Dieter’s first faculty position was at Occidental College in Eagle Rock, California, which he held for two years before accepting a professorship at Colorado State University in 1973, a position that he held for twenty years. During his Colorado years Dieter became known as an expert on the flora of the Rocky Mountains. It was also during this time that he married fellow botanist Beth Painter. Then, in 1991, Dieter was recruited to take over the management of The Jepson Manual Project. Dieter moved to UC Berkeley, where his combination of botanical acumen, ability to secure funding, and his ability to deal with the varied personalities of the authors shined. He also authored numerous treatments in the Manual (by my count he was the author or coordinator for ten families, plus author or co-author for another 46 genera), and he remains on the editorial board of The Jepson Manual Project today. After the first edition of The Jepson Manual was published in 1993, Dieter moved to his current home at the Santa Barbara Botanic Garden (SBBG) where he is Director of Conservation. Although Dieter studied the Asteraceae during his graduate years, he soon became interested in Polem- oniaceae, no doubt influenced by his major professor. He eventually published on Jpomopsis, Collomia, Allophyl- lum, and Polemonium. Today his Polemoniologist brethren (and sistren) consider him the reigning dean of the family. However, Dieter’s interests are not limited to Polemoniaceae. As just one example, he co-authored with Dave Fross the 2006 book Ceanothus, a monograph of the genus. Dieter’s research record (he published his first paper in 1967) has not only spanned an array of taxa, but has included floristics, taxonomic revisions, descriptions of new taxa, cytotaxonomy, palynology, chemosystematics, reproductive biology, and detailed monographic studies. In his role as Director of Conservation, he has also contributed valuable technical reports to various agen- cies. He is by all accounts a taxonomist’s taxonomist. Today Dtieter’s professional responsibilities include managing the SBBG conservation program, including acquisition and curation of ex situ plant materials; acting as liaison to various agencies; research on rare-plant biology; assisting the horticulture program on issues related to listed species; and managing the conservation collection and herbarium database. Dieter’s colleague at SBBG, Dr. Robert Muller, commented on Dieter’s other Important role at the Garden in “interpreting the complexities of botanical science for the interested layperson. He 1s a regular contributor to docent training and plays a critical role in providing background information to the education and horticulture programs. Of course, anyone who knows him can imagine the enthusiasm that he brings to it.” I first met Dieter when I was a young botany under- grad at UCSB and he was a (slightly older) graduate teaching assistant in Bob Haller’s California Flora course. This energetic botanist, who commonly wore shorts that we dubbed “‘Dieterhosen,” seemed to know the name of every plant, and it soon became clear that the students in Dieter’s lab sections had a leg up on the other students in the course. Several years later, when I became Dieter’s fellow lab mate in graduate school, the lab members would often go on weekend collecting trips. Too often, the rest of us rarely keyed anything; we simply went through our pressed material, and Dieter would tell us the names of every species (another lab member had christened him the ‘‘Walk-a-Munz’’). Dieter himself was not so lazy; he learned the plants by keying everything he saw. Anyone who has met Dieter knows that he is a most affable fellow, and given to tell stories in his own inimit- able style. (He does a great impression of the emi- nent botanist Ledyard Stebbins describing chromosome 240 behavior during meiosis.) In grad school we would delight as Dieter would tell us a joke or describe a movie he just saw. These stories are often punctuated with what have been called ‘‘Dieterisms,” similar in style to quotes made famous by Yogi Berra and Sam Goldwyn. A dedication such as this would not be complete without a few. Once in a seminar Dieter told us that “‘the snow persists until it melts.” Hmm ... How about, “this happens 99 times out of ten.’ Or, “I just noticed something that I’ve always known.” But the most famous Dieterism comes from a letter to me in 1983, whereupon having found some new species in the Rockies, he wrote “I am continually amazed at the things that have yet to be discovered.” The following year the American Association of Plant Taxonomists meeting was held at Colorado State University, and Dieter was our local host. When he was formally thanked at the annual dinner, Dave Seigler presented Dieter with a poster of the “continually amazed”’ quote printed on parchment. After the laughter and hooting subsided, Seigler announced that if anyone was interested in getting a copy, he had a few extras (he MADRONO [Vol. 59 pulled out a large stack) that he’d sell for 25 cents each. The crowd rushed to the podium to buy the posters. Then Art Cronquist asked Dieter to sign his copy. Dieter obliged, and a line instantly formed, with a large contingent of the American Society of Plant Taxono- mists waiting for Dr. Wilken’s signature. I still have my copy in my Office. In this year when the second edition of the Jepson Manual has been published, it 1s fitting that we recognize Dieter’s contribution as the principal editor of the first edition. Through his guidance and characteristic dili- gence, that volume was produced nearly 20 years ago and has served several generations of California botanists. So here’s to you, Dieter, for all of your contributions to the study of the California Flora, and for your valued collegiality to the botanical community. Bob Patterson Professor of Biology San Francisco State University San Francisco, CA MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME LIX 2002 BOARD OF EDITORS Class of: 2012—GRETCHEN LEBUHN, San Francisco State University, CA ROBERT PATTERSON, San Francisco State University, CA 2013—-ERIC ROALSON, Washington State University, WA KRISTINA SCHIERENBECK, California State University, Chico, CA 2014—BRANDON PRATT, California State University, Bakersfield, CA TOM WENDT, University of Texas, Austin, TX Corresponding Editor—MATT RITTER Biological Sciences Department Cal Poly, San Luis Obispo 1 Grand Avenue San Luis Obispo, CA 93407 madronoeditor@gmail.com AND Copy Editor—RICHARD WHITKUS Department of Biology Sonoma State University 1801 E. Cotati Avenue Rohnert Park, CA 94928-3609 whitkus@sonoma.edu Published quarterly by the California Botanical Society, Inc. Life Sciences Building, University of California, Berkeley 94720 Printed by Allen Press, Inc., Lawrence, KS 66044 MADRONO VOLUME 59 TABLE OF CONTENTS Aho, Ken, and Janet Bala, Vascular alpine flora of Mount Washburn, Yellowstone National Park, USA —__ Alexander, Earl B., Comment on the gabbro soils of Pine Hill Allen, Edith B. (see Steers, Robert J.) Bala, Janet (see Aho, Ken) Bell, Duncan (see Fraga, Naomi S.) Benedict, Beverly G., et al., Mimulus sookensis (Phrymaceae), a new allotetraploid species derived from Mimulus guttatus and Mimulus nasutus Bowen, Lizabeth (see Van Vuren, Dirk H.) Brummitt, R. K., Scott D. White and Justin M. Wood, Status of Bingham’s morning-glory in the light of its TESCO VSI) Secs ss ee ee ne se eae ee ee ee Canter, Adam N.., Noteworthy collection from California. eee Crow, Taylor, and Matt Ritter, Point of View; Changes to the botanical code and what they mean for western ANOrihnsAmerncan: DOlANY” 222.05 8S A ee ee ee Culley, Theresa M., and Richard L. Stokes, Genetic structure and outcrossing rates in Viola pedunculata (Violaceae), a California endemic violet lacking cleistogamous flowers DeWoody, Jennifer, Valerie D. Hipkins, Julie Kierstead Nelson and Len Lindstrand, III, Genetic structure of Vaccinium parvifolium (Ericaceae) in northern California reveals potential systematic distinctions __ Dodero, Mark W., and Michael G. Simpson, Dudley crassifolia (Crassulaceae), a new species from northern Baja California, Mexico Ellison, Nicholas (see Heise, Kerry) Fotheringham, C. J. (see Keeley, Jon E.) Fraga, Naomi S., Elizabeth Kempton, LeRoy Gross and Duncan Bell, Reappearance of the vanishing wild buckwheat: a status review of Eriogonum evanidum (Polygonaceae) | Ganders, Fred R. (see Benedict, Beverly G., et al.) Gross, LeRoy (see Fraga, Naomi S.) Guilliams, C. Matt, Review of Research and Discovery in Vernal Pool Landscapes, edited by D. G. Alexander and R. A. Schlising Iss a ee mee at Pace een ere see Hannan, Gary L., Change in rank of Er iodicty on traskiae subsp. smithii (Hydrophyllaceae) Ano ee Heise, Kerry, Geri Hulse-Stephens, and Nicholas Ellison, Noteworthy collection from California Hipkins, Valerie D. (see Dewoody, Jennifer) Hrusa, G. F., Morphological and isoenzyme variation in Rhododendron occidentale (western azalea) (section Pentathera; Ericaceae) Hulse-Stephens, Geri (see Heise, Kerry) Keeley, Jon E., C. J. Fotheringham and Philip W. Rundel, Postfire chaparral regeneration under Mediterranean and non-mediterranean climates Kempton, Elizabeth (see Fraga, Naomi S.) Kelch, Dean G., and Andrew Murdock, Flora of the Carquinez Strait region, Contra Costa and Solano Counties, California ee. Les, Donald, Nicholas Tippery and Hamid Razifard, Noteworthy collection from California _ Les, Donald, Nicholas Tippery and Hamid Razifard, Noteworthy collection from Texas Lindstrand, Len, III (see Dewoody, Jennifer) Martin, Noland H. (see Benedict, Beverly G., et al.) Modliszewski, Jennifer L. (see Benedict, Beverly G., et al.) Murdock, Andrew (see Kelch, Dean G.) Nelson, Julie Kierstead (see Dewoody, Jennifer) Odegard, Craig, Noteworthy collection from Montana Parker, V. Thomas, President’s report for Volume 59 Patterson, Bob, Dedication of Volume 59 to Dieter H. Wilken | Patterson, Robert (see Schultz, Joanna L.) Patterson, Robert (see also Walden, Genevieve K.) Razifard, Hamid (see Les, Donald - two articles) Riegel, Gregg M. (see Weixelman, Dave A.) Ritter, Matt (see Crow, Taylor) Ritter, Matt, and Richard Whitkus, Editors’ Report for Volume 59 Rundel, Philip W. (see Keeley, Jon E.) Sarr, Daniel A., Review of Northwest California: A Natural History by John O. Sawyer Schultz, Joanna L., and Robert Patterson, A new combination in Linanthus (Polemoniaceae) from Idaho and Oregon Shevock, James R. (see Spence, John R.) Simpson, Michael G. (see Dodero, Mark W.) 150 [Vol. 59 TABLE OF CONTENTS Spence, John R., and James R. Shevock, Ptychostomum pacificum (Bryaceae), a new fen species from California, Oregon and western Nevada, USA Steers, Robert J., and Edith B. Allen, Impact of recurrent fire on annual plants: A case study from the western edge of the Colorado Desert *. ere Es Stokes, Richard L. (see Culley, Theresa M.) Sweigart, Andrea L. (see Benedict, Beverly G., et al.) Tippery, Nicholas (see Les, Donald — two articles) Van Vuren, Dirk H., and Lizabeth Bowen, Response of grassland vegetation on Santa Cruz Island to removal of feral sheep Walden, Genevieve K., and Robert Patterson, Nomenclature of subdivisions within Phacelia (Boraginaceae: Hydrophylloideae) - Weixelman, Dave A., and Gregg M. ‘Riegel, “Measurement of ‘spatial autocorrelation of vegetation in mountain meadows of the Sierra Nevada, California and western Nevada White, Scott D. (see Brummitt, R. K.) Whitkus, Richard (see Ritter, Matt, and Richard Whitkus) Willis, John H. (see Benedict, Beverly G., et al.) Wilson, Barbara L. (see Zika, Peter F.) Wood, Justin M. (see Brummitt, R. K.) York, Dana A., Noteworthy collection from California Zika, Peter F., and Barbara L. Wilson, Carex albida (Cyperaceae), and its relationship to Carex lemmonii ll 171 DATES OF PUBLICATION OF MADRONO, VOLUME 59 Number |, pages 1-46, published 6 July 2012 Number 2, pages 47-108, published 27 August 2012 Number 3, pages 109-168, published 27 August 2012 Number 4, pages 169—240, published 19 December 2012 SUBSCRIPTIONS — MEMBERSHIP The California Botanical Society has several membership types (individuals ($40 per year; family $45 per year; emeritus $32 per year; students $27 per year for a maximum of 7 years). Late fees may be assessed. Beginning in 2011, rates will increase by $5 for all membership types except life memberships, for which rates will increase by $100, and student memberships, which will not show a rate increase. Members of the Society receive MADRONO free. Institutional subscriptions to MADRONO are available ($75). Membership is based on a calendar year only. Life memberships are $850. Applications for membership (including dues), orders for subscriptions, and renewal payments should be sent to the Membership Chair. Requests and rates for back issues, changes of address, and undelivered copies of MADRONO should be sent to the Corresponding Secretary. INFORMATION FOR CONTRIBUTORS Submit manuscripts to http://www.edmgr.com/madrono/. If you are submitting a manuscript for the first time to the MADRONO PeerTrack website, you must first register by clicking "register now" and following the instructions. Manuscripts by authors having outstanding page charges will not be sent for review. Manuscripts may be submitted in English or Spanish. English-language manuscripts dealing with taxa or topics of Latin America and Spanish-language manuscripts must have a Spanish RESUMEN and an English ABSTRACT. For all articles and short items (NOTES, NOTEWORTHY COLLECTIONS, POINTS OF VIEW, etc.), follow the format used in recent issues for the type of item submitted. Detailed information for contributors, including instructions for manuscript preparation, is available at the California Botanical Society website, www.calbotsoc. org; follow the link for MADRONo. Authors are encouraged to include the names, addresses, and e-mail addresses of two to four potential reviewers with their submitted manuscript. All California Botanical Society members current in the volume year that their contributions are published are allowed five free pages per volume year. Additional pages will be charged at the rate of $40 per page. Joint authors may apply all or a portion of their respective five-page allotments to a jointly-published article. Partial pages will be charged as full. The purpose of this fee is not to pay directly for the costs of publishing any particular paper, but rather to allow the Society to continue publishing MADRONO on a reasonable schedule, with equity among all members for access to its pages. Printer’s fees for color plates and other complex matter (including illustrations, charts, maps, photographs) will be charged at cost. Author’s changes after typesetting @ $4.50 per line will be charged to authors. Page charges are important in maintaining Madrono as a viable publication, and timely payment of charges is appreciated. At the time of submission, authors must provide information describing the extent to which data in the manuscript have been used in other papers that are published, in press, submitted, or soon to be submitted elsewhere. wi 3 9088 01700 3 ANNOUNCEMENTS ~_—)sPresiwent ’s REPORT FOR VOLUME 59 EDITORS’ Vo REVIEWERS OR ee i a AMC CAN) ae EDICATION oesscccessssseeecstteessseeesentssssseeeesttnnnssteeeestnsnsseeesssnsssssesssstsssssseessises 234 sisldlcis ele a le/bioleielg|s/sls wicictsleistelere etm